ArticleTitle stringclasses 109 values | Question stringlengths 4 586 ⌀ | Answer stringlengths 1 926 ⌀ | ArticleFile stringclasses 57 values | EvidencesAvailable stringclasses 120 values |
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Amedeo_Avogadro | Why did Avogadro lose his chair at the University of Turin? | He was active in the revolutionary movements of 1821 against the king of Sardinia | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Why did Avogadro lose his chair at the University of Turin? | Avogadro lost his chair at the University of Turin because he was active in the revolutionary movements of 1821 against the king of Sardinia. | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Was Amedeo Avogadro was born in Turin? | Yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Was Amedeo Avogadro was born in Turin? | yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Is Avogadro hailed as a founder of the atomic-molecular theory? | Yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Is Avogadro hailed as a founder of the atomic-molecular theory? | yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Did Johann Josef Loschmidt first calculate the value of Avogadro's number? | Yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Did Johann Josef Loschmidt first calculate the value of Avogadro's number? | yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | In 1820, Avogadro became a professor of physics where? | University of Turin | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | In 1820, Avogadro became a professor of physics where? | University of Turin | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Avogadro did not actually use what word? | atom | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Avogadro did not actually use what word? | atom | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | The number of elementary entities in 1 mole of a substance is known as what? | Avogadro constant | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | The number of elementary entities in 1 mole of a substance is known as what? | Avogadro's constant | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | What would a German call Avogadro's number? | Loschmidt number | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | What would a German call Avogadro's number? | Loschmidt number | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Is Amedeo Avogadro Italian? | Yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Is Amedeo Avogadro Italian? | yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Did he become a professor before the revolutionary movements against the king of Sardinia? | Yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Amedeo_Avogadro | Did he become a professor before the revolutionary movements against the king of Sardinia? | yes | data/set4/a8 | Amedeo_Avogadro
Lorenzo Romano Amedeo Carlo Avogadro di Quaregna (Quaregga) e di Cerreto, Count of Quaregna (or Quaregga) and Cerreto (9 August 1776 â 9 July 1856) was an Italian savant. He is most noted for his contributions to molecular theory, including what is known as Avogadro's law. In tribute to him, the number of elementary entities (atoms, molecules, ions or other particles) in 1 mole of a substance, , is known as the Avogadro constant.
Amedeo Avogadro was born in Turin to a noble family of Piedmont, Italy.
He graduated in ecclesiastical law at the early age of 20 and began to practice. Soon after, he dedicated himself to physics and mathematics (then called positive philosophy), and in 1809 started teaching them at a liceo (high school) in Vercelli, where his family had property.
In 1811, he published an article with the title Essai d'une manière de déterminer les masses relatives des molécules élémentaires des corps, et les proportions selon lesquelles elles entrent dans ces combinaisons ("Essay on Determining the Relative Masses of the Elementary Molecules of Bodies and the Proportions by Which They Enter These Combinations"), which contains Avogadro's hypothesis. Avogadro submitted this essay to a French journal, De Lamétherie's Journal de Physique, de Chimie et d'Histoire naturelle (Journal of Physics, Chemistry and Natural History) so it was written in French, not Italian. (Note: In 1811, northern Italy was under the rule of the French Emperor Napoléon Bonaparte.)
In 1820, he became professor of physics at the University of Turin. After the downfall of Napoléon in 1815, northern Italy came under control of this kingdom.
He was active in the revolutionary movements of 1821 against the king of Sardinia (who became ruler of Piedmont with Turin as his capital). As a result, he lost his chair in 1823 (or the university officially declared, it was "very glad to allow this interesting scientist to take a rest from heavy teaching duties, in order to be able to give better attention to his researches") .
Eventually, Charles Albert granted a Constitution (Statuto Albertino) in 1848. Well before this, Avogadro had been recalled to the university in Turin in 1833, where he taught for another twenty years.
Little is known about Avogadro's private life, which appears to have been sober and religious. He married Felicita Mazzé and had six children.
Some historians suggest that he sponsored some Sardinian revolutionaries, who were stopped by the announcement of Charles Albert's constitution.
Avogadro held posts dealing with statistics, meteorology, and weights and measures (he introduced the metric system into Piedmont) and was a member of the Royal Superior Council on Public Instruction.
In honor of Avogadro's contributions to molecular theory, the number of molecules in one mole was named Avogadro's number, N A or "Avogadro's constant". It is approximately 6.0221415 10 23 . Avogadro's number is used to compute the results of chemical reactions. It allows chemists to determine the exact amounts of substances produced in a given reaction.
Johann Josef Loschmidt first calculated the value of Avogadro's number, often referred to as the Loschmidt number in German-speaking countries (Loschmidt constant now has another meaning).
Avogadro's Law states that the relationship between the masses of the same volume of different gases (at the same temperature and pressure) corresponds to the relationship between their respective molecular weights. Hence, the relative molecular mass of a gas can be calculated from the mass of sample of known volume.
Avogadro developed this hypothesis after Joseph Louis Gay-Lussac had published in 1808 the Gay-Lussac law his law on volumes (and combining gases). The greatest problem Avogadro had to resolve was the confusion at that time regarding atoms and molecules. One of his most important contributions was clearly distinguishing one from the other, stating that gases are composed of molecules, and these molecules are composed of atoms. For instance, John Dalton did not consider this possibility. Avogadro did not actually use the word "atom" as the words "atom" and "molecule" were used almost without difference. He believed that there were three kinds of "molecules," including an "elementary molecule" (our "atom"). Also, more attention was given to the definition of mass, as distinguished from weight.
In 1814, he published Mémoire sur les masses relatives des molécules des corps simples, ou densités présumées de leur gaz, et sur la constitution de quelques-uns de leur composés, pour servir de suite à l'Essai sur le même sujet, publié dans le Journal de Physique, juillet 1811 ("Note on the Relative Masses of Elementary Molecules, or Suggested Densities of Their Gases, and on the Constituents of Some of Their Compounds, As a Follow-up to the Essay on the Same Subject, Published in the Journal of Physics, July 1811") (), about gas densities.
In 1821 he published another paper, Nouvelles considérations sur la théorie des proportions déterminées dans les combinaisons, et sur la détermination des masses des molécules des corps (New Considerations on the Theory of Proportions Determined in Combinations, and on Determination of the Masses of Atoms) and shortly afterwards, Mémoire sur la manière de ramener les composès organiques aux lois ordinaires des proportions déterminées (Note on the Manner of Finding the Organic Composition by the Ordinary Laws of Determined Proportions).
In 1841, he published his work in Fisica dei corpi ponderabili, ossia Trattato della costituzione materiale de' corpi, 4 volumes.
The scientific community did not give great attention to his theory, so Avogadro's hypothesis was not immediately accepted. André-Marie Ampère achieved the same results three years later by another method (in his -- On the Determination of Proportions in which Bodies Combine According to the Number and the Respective Disposition of the Molecules by Which Their Integral Particles Are Made), but the same indifference was shown to his theory as well.
Only through studies by Charles Frédéric Gerhardt and Auguste Laurent on organic chemistry was it possible to demonstrate that Avogadro's law explained why the same quantities of molecules in a gas have the same volume.
Unfortunately, related experiments with some inorganic substances showed seeming exceptions to the law. This was finally resolved by Stanislao Cannizzaro, as announced at Karlsruhe Congress in 1860, four years after Avogadro's death. He explained that these exceptions were due to molecular dissociations at certain temperatures, and that Avogadro's law determined not only molecular masses, but atomic masses as well.
In 1911, a meeting in Turin commemorated the hundredth anniversary of the publication of Avogadro's classic 1811 paper. King Victor Emmanuel III attended. Thus, Avogadro's great contribution to chemistry was recognized.
Rudolf Clausius, with his kinetic theory on gases, gave another confirmation of Avogadro's Law. Jacobus Henricus van 't Hoff showed that Avogadro's theory also held in dilute solutions.
Avogadro is hailed as a founder of the atomic-molecular theory.
*
*
*
* Morselli, Mario. (1984). Amedeo Avogadro, a Scientific Biography. Kluwer. ISBN 9027716242.
:*Review of Morselli's book:
* Avogadro (lunar crater)
* Avogadro's constant
|
Ant | Do ants belong to the Hymenoptera order? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do ants belong to the Hymenoptera order? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are ants used in cuisine? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are ants used in cuisine? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Does an ant's head contain sensory organs? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Does an ant's head contain sensory organs? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | How do most ants travel? | most ants travel by walking | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | How do most ants travel? | by walking | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | In ant colonies, what are the fertile female ants called? | queens | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | In ant colonies, what are the fertile female ants called? | queens | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Who wrote about ants in A Tramp Abroad? | mark twain wrote about ants | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Who wrote about ants in A Tramp Abroad? | Mark Twain | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do the ants eat plants, meats, or both? | both | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What organs gives a bull ant its good sight? | compound eyes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What may happen to red fire ants if we use boiling water on the queen? | nests of red fire ants may be destroyed | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What may happen to red fire ants if we use boiling water on the queen? | die | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are ants found in Antartica? | No, there are no ants in antarctica. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are ants found in Antartica? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do male ants take flight before females? | Yes, male ants take flight before females. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do male ants take flight before females? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do worker ants have wings? | No, worker ants do not have wings. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do worker ants have wings? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What are the three segments of an ant? | The head, mesosoma and metasoma are the three distinct body segments. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What are the three segments of an ant? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What are the ant colonies that lack queens called? | Colonies that lack queens are called gamergate colonies. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What are the ant colonies that lack queens called? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Where are bullet ants located? | Bullet ants are located in Central and South America. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Where are bullet ants located? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do ants belong to the same order as bees? | Yes, ants belong to the same order as bees. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do ants belong to the same order as bees? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What is the most common color of ants? | Most ants are red or black. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | What is the most common color of ants? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do all ants build nests? | No, not all ants build nests. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do all ants build nests? | null | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Ultraviolet vision was first discovered in ants by Sir who in 1881? | John Lubbok | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do ants thrive in most ecosystems? | Yes. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are several species of ants able to use the Earths magnetic field? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are termites actually more closely related to cockroaches as well as mantids? | Yes, termites are actually more closely related to cockroaches as well as mantids. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Do some caterpillars produce vibrations as well as sounds? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Did ants evolve from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants? | yes | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Is the ant a marsupial? | no | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Ant | Are they easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist? | Yes, they are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist. | data/set1/a1 | Ant
Ants are social insects of the family Formicidae ( ), and along with the related wasps and bees, they belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. Today, more than 12,500 species are classified with upper estimates of about 22,000 species. They are easily identified by their elbowed antennae and a distinctive node-like structure that forms a slender waist.
Ants form colonies that range in size from a few dozen predatory individuals living in small natural cavities to highly organised colonies which may occupy large territories and consist of millions of individuals. These larger colonies consist mostly of sterile wingless females forming castes of "workers", "soldiers", or other specialised groups. Nearly all ant colonies also have some fertile males called "drones" and one or more fertile females called "queens". The colonies are sometimes described as superorganisms because the ants appear to operate as a unified entity, collectively working together to support the colony.
Ants have colonised almost every landmass on Earth. The only places lacking indigenous ants are Antarctica and certain remote or inhospitable islands. Ants thrive in most ecosystems, and may form 15 25% of the terrestrial animal biomass. Their success has been attributed to their social organisation and their ability to modify habitats, tap resources, and defend themselves. Their long co-evolution with other species has led to mimetic, commensal, parasitic, and mutualistic relationships. Hölldobler & Wilson (1990), p. 471
Ant societies have division of labour, communication between individuals, and an ability to solve complex problems. These parallels with human societies have long been an inspiration and subject of study.
Many human cultures make use of ants in cuisine, medication and rituals. Some species are valued in their role as biological pest control agents. However, their ability to exploit resources brings ants into conflict with humans, as they can damage crops and invade buildings. Some species, such as the red imported fire ant, are regarded as invasive species, since they have established themselves in new areas where they have been accidentally introduced.
The word ant is derived from ante of Middle English which is derived from æmette of Old English and is related to the Old High German Ämeiza, hence the modern German Ameise. All of these words come from West Germanic *amaitjo, and the original meaning of the word was "the biter" (from Proto-Germanic *ai-, "off, away" + *mait- "cut"). The family name Formicidae is derived from the Latin formÄ«ca ("ant") from which the words in other Romance languages such as the Portuguese formiga, Italian formica, Spanish hormiga, Romanian furnicÄ and French fourmi are derived.
Ants fossilised in Baltic amber
The family Formicidae belongs to the order Hymenoptera, which also includes sawflies, bees and wasps. Ants evolved from a lineage within the vespoid wasps. Phylogenetic analysis suggests that ants arose in the mid-Cretaceous period about 110 to 130 million years ago. After the rise of flowering plants about 100 million years ago they diversified and assumed ecological dominance around 60 million years ago.
In 1966, E. O. Wilson and his colleagues identified the fossil remains of an ant (Sphecomyrma freyi) that lived in the Cretaceous period. The specimen, trapped in amber dating back to more than 80 million years ago, has features of both ants and wasps. Sphecomyrma was probably a ground forager but some suggest on the basis of groups such as the Leptanillinae and Martialinae that primitive ants were likely to have been predators under the soil surface.
Phylogenetic position of the Formicidae.
During the Cretaceous period, a few species of primitive ants ranged widely on the Laurasian super-continent (the northern hemisphere). They were scarce in comparison to other insects, representing about 1% of the insect population. Ants became dominant after adaptive radiation at the beginning of the Tertiary period. By the Oligocene and Miocene ants had come to represent 20 40% of all insects found in major fossil deposits. Of the species that lived in the Eocene epoch, approximately one in ten genera survive to the present. Genera surviving today comprise 56% of the genera in Baltic amber fossils (early Oligocene), and 92% of the genera in Dominican amber fossils (apparently early Miocene). Hölldobler & Wilson (1990), pp. 23 24
Termites, though sometimes called white ants, are not ants and belong to the order Isoptera. Termites are actually more closely related to cockroaches and mantids. Termites are eusocial but differ greatly in the genetics of reproduction. The similar social structure is attributed to convergent evolution. Velvet ants look like large ants, but are wingless female wasps.
Ants are found on all continents except Antarctica and only a few large islands such as Greenland, Iceland, parts of Polynesia and the Hawaiian Islands lack native ant species. Ants occupy a wide range of ecological niches, and are able to exploit a wide range of food resources either as direct or indirect herbivores, predators and scavengers. Most species are omnivorous generalists but a few are specialist feeders. Their ecological dominance may be measured by their biomass, and estimates in different environments suggest that they contribute 15 20% (on average and nearly 25% in the tropics) of the total terrestrial animal biomass, which exceeds that of the vertebrates.
Ants range in size from . Hölldobler & Wilson (1990), p. 589 Their colours vary; most are red or black, green is less common, and some tropical species have a metallic lustre. More than 12,000 species are currently known (with upper estimates of about 14,000), with the greatest diversity in the tropics. Taxonomic studies continue to resolve the classification and systematics of ants. Online databases of ant species, including AntBase and the Hymenoptera Name Server, help to keep track of the known and newly described species. The relative ease with which ants can be sampled and studied in ecosystems has made them useful as indicator species in biodiversity studies.
Ants are distinct in their morphology from other insects in having elbowed antennae, metapleural glands, and a strong constriction of their second abdominal segment into a node-like petiole. The head, mesosoma and metasoma or gaster are the three distinct body segments. The petiole forms a narrow waist between their mesosoma (thorax plus the first abdominal segment, which is fused to it) and gaster (abdomen less the abdominal segments in the petiole). The petiole can be formed by one or two nodes (the second alone, or the second and third abdominal segments). Borror, Triplehorn & Delong (1989), p. 737
Bull ant showing the powerful mandibles and the relatively large compound eyes that provide excellent vision
Like other insects, ants have an exoskeleton, an external covering that provides a protective casing around the body and a point of attachment for muscles, in contrast to the internal skeletons of humans and other vertebrates. Insects do not have lungs; oxygen and other gases like carbon dioxide pass through their exoskeleton through tiny valves called spiracles. Insects also lack closed blood vessels; instead, they have a long, thin, perforated tube along the top of the body (called the "dorsal aorta") that functions like a heart, and pumps haemolymph towards the head, thus driving the circulation of the internal fluids. The nervous system consists of a ventral nerve cord that runs the length of the body, with several ganglia and branches along the way reaching into the extremities of the appendages. Borror, Triplehorn & Delong (1989), pp. 24 71
Diagram of a worker ant (Pachycondyla verenae)
An ant's head contains many sensory organs. Like most insects, ants have compound eyes made from numerous tiny lenses attached together. Ants' eyes are good for acute movement detection but do not give a high resolution. They also have three small ocelli (simple eyes) on the top of the head that detect light levels and polarization. Compared to vertebrates, most ants have poor-to-mediocre eyesight and a few subterranean species are completely blind. Some ants such as Australia's bulldog ant, however, have exceptional vision. Two antennae ("feelers") are attached to the head; these organs detect chemicals, air currents and vibrations; they are also used to transmit and receive signals through touch. The head has two strong jaws, the mandibles, used to carry food, manipulate objects, construct nests, and for defence. In some species a small pocket (infrabuccal chamber) inside the mouth stores food, so it can be passed to other ants or their larvae.
All six legs are attached to the mesosoma ("thorax"). A hooked claw at the end of each leg helps ants to climb and hang onto surfaces. Most queens and male ants have wings; queens shed the wings after the nuptial flight, leaving visible stubs, a distinguishing feature of queens. However, wingless queens (ergatoids) and males occur in a few species.
The metasoma (the "abdomen") of the ant houses important internal organs, including those of the reproductive, respiratory (tracheae) and excretory systems. Workers of many species have their egg-laying structures modified into stings that are used for subduing prey and defending their nests.
Seven Leafcutter ant workers of various castes (left) and two Queens (right)
In the colonies of a few ant species, there are physical castes workers in distinct size-classes, called minor, median, and major workers. Often the larger ants have disproportionately larger heads, and correspondingly stronger mandibles. Such individuals are sometimes called "soldier" ants because their stronger mandibles make them more effective in fighting, although they are still workers and their "duties" typically do not vary greatly from the minor or median workers. In a few species the median workers are absent, creating a sharp divide between the minors and majors. Weaver ants, for example, have a distinct bimodal size distribution.
Some other species show continuous variation in the size of workers. The smallest and largest workers in Pheidologeton diversus show nearly a 500-fold difference in their dry-weights. Workers cannot mate; however, because of the haplodiploid sex-determination system in ants, workers of a number of species can lay unfertilised eggs that become fully fertile haploid males. The role of workers may change with their age and in some species, such as honeypot ants, young workers are fed until their gasters are distended, and act as living food storage vessels. These food storage workers are called repletes. This polymorphism in morphology and behaviour of workers was initially thought to be determined by environmental factors such as nutrition and hormones which led to different developmental paths; however, genetic differences between worker castes have been noted in Acromyrmex sp. These polymorphisms are caused by relatively small genetic changes; differences in a single gene of Solenopsis invicta can decide whether the colony will have single or multiple queens. The Australian jack jumper ant (Myrmecia pilosula), has only a single pair of chromosomes (males have just one chromosome as they are haploid), the lowest number known for any animal, making it an interesting subject for studies in the genetics and developmental biology of social insects.
Meat eater ant nest during swarming
The life of an ant starts from an egg. If the egg is fertilised, the progeny will be female (diploid); if not, it will be male (haploid). Ants develop by complete metamorphosis with the larval stages passing through a pupal stage before emerging as an adult. The larva is largely immobile and is fed and cared for by workers. Food is given to the larvae by trophallaxis, a process in which an ant regurgitates liquid food held in its crop. This is also how adults share food, stored in the "social stomach", among themselves. Larvae may also be provided with solid food such as trophic eggs, pieces of prey and seeds brought back by foraging workers and may even be transported directly to captured prey in some species. The larvae grow through a series of moults and enter the pupal stage. The pupa has the appendages free and not fused to the body as in a butterfly pupa. The differentiation into queens and workers (which are both female), and different castes of workers (when they exist), is determined by the nutrition the larvae obtain. Larvae and pupae need to be kept at fairly constant temperatures to ensure proper development, and so are often moved around the various brood chambers within the colony. Hölldobler & Wilson (1990), pp. 351, 372
A new worker spends the first few days of its adult life caring for the queen and young. It then graduates to digging and other nest work, and later to defending the nest and foraging. These changes are sometimes fairly sudden, and define what are called temporal castes. An explanation for the sequence is suggested by the high casualties involved in foraging, making it an acceptable risk only for ants that are older and are likely to die soon of natural causes.
Fertilised meat eater ant queen beginning to dig a new colony
Most ant species have a system in which only the queen and breeding females have the ability to mate. Contrary to popular belief, some ant nests have multiple queens while others can exist without queens. Workers with the ability to reproduce are called "gamergates" and colonies that lack queens are then called gamergate colonies; colonies with queens are said to be queen-right. The winged male ants, called drones, emerge from pupae along with the breeding females (although some species, like army ants, have wingless queens), and do nothing in life except eat and mate. Most ants are univoltine, producing a new generation each year. During the species specific breeding period, new reproductives, winged males and females leave the colony in what is called a nuptial flight. Typically, the males take flight before the females. Males then use visual cues to find a common mating ground, for example, a landmark such as a pine tree to which other males in the area converge. Males secrete a mating pheromone that females follow. Females of some species mate with just one male, but in some others they may mate with anywhere from one to ten or more different males. Hölldobler & Wilson (1990) Mated females then seek a suitable place to begin a colony. There, they break off their wings and begin to lay and care for eggs. The females store the sperm they obtain during their nuptial flight to selectively fertilise future eggs. The first workers to hatch are weak and smaller than later workers, but they begin to serve the colony immediately. They enlarge the nest, forage for food and care for the other eggs. This is how new colonies start in most species. Species that have multiple queens may have a queen leaving the nest along with some workers to found a colony at a new site, Hölldobler & Wilson (1990), pp. 143 179 a process akin to swarming in honeybees.
Ants mating.
A wide range of reproductive strategies have been noted in ant species. Females of many species are known to be capable of reproducing asexually through thelytokous parthenogenesis and one species, Mycocepurus smithii is known to be all-female.
Ant colonies can be long-lived. The queens can live for up to 30 years, and workers live from 1 to 3 years. Males, however, are more transitory, and survive only a few weeks. Ant queens are estimated to live 100 times longer than solitary insects of a similar size.
Ants are active all year long in the tropics but, in cooler regions, survive the winter in a state of dormancy or inactivity. The forms of inactivity are varied and some temperate species have larvae going into the inactive state (diapause), while in others, the adults alone pass the winter in a state of reduced activity.
Weaver ants collaborating to dismember a red ant (the two at the extremities are pulling the red ant, while the middle one cuts the red ant until it snaps)
Ants communicate with each other using pheromones. These chemical signals are more developed in ants than in other hymenopteran groups. Like other insects, ants perceive smells with their long, thin and mobile antennae. The paired antennae provide information about the direction and intensity of scents. Since most ants live on the ground, they use the soil surface to leave pheromone trails that can be followed by other ants. In species that forage in groups, a forager that finds food marks a trail on the way back to the colony; this trail is followed by other ants, these ants then reinforce the trail when they head back with food to the colony. When the food source is exhausted, no new trails are marked by returning ants and the scent slowly dissipates. This behaviour helps ants deal with changes in their environment. For instance, when an established path to a food source is blocked by an obstacle, the foragers leave the path to explore new routes. If an ant is successful, it leaves a new trail marking the shortest route on its return. Successful trails are followed by more ants, reinforcing better routes and gradually finding the best path.
Ants use pheromones for more than just making trails. A crushed ant emits an alarm pheromone that sends nearby ants into an attack frenzy and attracts more ants from further away. Several ant species even use "propaganda pheromones" to confuse enemy ants and make them fight among themselves. Pheromones are produced by a wide range of structures including Dufour's glands, poison glands and glands on the hindgut, pygidium, rectum, sternum and hind tibia. Pheromones are also exchanged mixed with food and passed by trophallaxis, transferring information within the colony. This allows other ants to detect what task group (e.g., foraging or nest maintenance) other colony members belong to. In ant species with queen castes, workers begin to raise new queens in the colony when the dominant queen stops producing a specific pheromone. Hölldobler & Wilson (1990), p. 354
Some ants produce sounds by stridulation, using the gaster segments and their mandibles. Sounds may be used to communicate with colony members or with other species.
A weaver ant in fighting position, mandibles wide open
Ants attack and defend themselves by biting and, in many species, by stinging, often injecting or spraying chemicals like formic acid. Bullet ants (Paraponera), located in Central and South America, are considered to have the most painful sting of any insect, although it is usually not fatal to humans. This sting is given the highest rating on the Schmidt Sting Pain Index. The sting of Jack jumper ants can be fatal, and an antivenin has been developed. Fire ants, Solenopsis spp., are unique in having a poison sac containing piperidine alkaloids. Their stings are painful and can be dangerous to hypersensitive people.
Trap-jaw ants of the genus Odontomachus are equipped with mandibles called trap-jaws, which snap shut faster than any other predatory appendages within the animal kingdom. One study of Odontomachus bauri recorded peak speeds of between 126 and 230 km/h (78 - 143 mph), with the jaws closing within 130 microseconds on average. The ants were also observed to use their jaws as a catapult to eject intruders or fling themselves backwards to escape a threat. Before the strike, the ant opens its mandibles extremely widely and locks them in this position by an internal mechanism. Energy is stored in a thick band of muscle and explosively released when triggered by the stimulation of sensory hairs on the inside of the mandibles. The mandibles also permit slow and fine movements for other tasks. Trap-jaws are also seen in the following genera: Anochetus, Orectognathus, and Strumigenys, plus some members of the Dacetini tribe , which are viewed as examples of convergent evolution. A Malaysian species of ant in the Camponotus cylindricus group has enlarged mandibular glands that extend into their gaster. When disturbed, workers rupture the membrane of the gaster, causing a burst of secretions containing acetophenones and other chemicals that immobilise small insect attackers. The worker subsequently dies. Suicidal defences by workers are also noted in a Brazilian ant Forelius pusillus where a small group of ants leaves the security of the nest after sealing the entrance from the outside each evening.
Ant mound holes prevent water from entering the nest during rain.
In addition to defence against predators, ants need to protect their colonies from pathogens. Some worker ants maintain the hygiene of the colony and their activities include undertaking or necrophory, the disposal of dead nest-mates. Oleic acid has been identified as the compound released from dead ants that triggers necrophoric behaviour in Atta mexicana while workers of Linepithema humile react to the absence of characteristic chemicals (dolichodial and iridomyrmecin) present on the cuticle of their living nestmates.
Nests may be protected from physical threats such as flooding and over-heating by elaborate nest architecture. Workers of Cataulacus muticus, an arboreal species that lives in plant hollows, respond to flooding by drinking water inside the nest, and excreting it outside. Camponotus anderseni which is nests in cavities within wood in mangrove habitats deals with submergence under water by switching to anaerobic respiration.
Many animals can learn behaviours by imitation but ants may be the only group apart from mammals where interactive teaching has been observed. A knowledgeable forager of Temnothorax albipennis leads a naive nest-mate to newly discovered food by the process of tandem running. The follower obtains knowledge through its leading tutor. Both leader and follower are acutely sensitive to the progress of their partner with the leader slowing down when the follower lags, and speeding up when the follower gets too close.
Controlled experiments with colonies of Cerapachys biroi suggest that individuals may choose nest roles based on their previous experience. An entire generation of identical workers was divided into two groups whose outcome in food foraging was controlled. One group was continually rewarded with prey, while it was made certain that the other failed. As a result, members of the successful group intensified their foraging attempts while the unsuccessful group ventured out less and less. A month later, the successful foragers continued in their role while the others moved to specialise in brood care.
Leaf nest of weaver ants, Pamalican, Philippines
Complex nests are built by many ants, but other species are nomadic and do not build permanent structures. Ants may form subterranean nests or build them on trees. These nests can be found in the ground, under stones or logs, inside logs, hollow stems or even acorns. The materials used for construction include soil and plant matter, and ants carefully select their nest sites; Temnothorax albipennis will avoid sites with dead ants, as these may indicate the presence of pests or disease. They are quick to abandon established nests at the first sign of threats.
The army ants of South America and the driver ants of Africa do not build permanent nests, but instead alternate between nomadism and stages where the workers form a temporary nest (bivouac) from their own bodies, by holding each other together. Hölldobler & Wilson (1990), p. 573
Weaver ant (Oecophylla spp.) workers build nests in trees by attaching leaves together, first pulling them together with bridges of workers and then inducing their larvae to produce silk as they are moved along the leaf edges. Similar forms of nest construction are seen in some species of Polyrhachis.
Myrmecocystus (Honeypot) ants store food to prevent colony famine.
Most ants are generalist predators, scavengers and indirect herbivores, but a few have evolved specialised ways of obtaining nutrition. Leafcutter ants (Atta and Acromyrmex) feed exclusively on a fungus that grows only within their colonies. They continually collect leaves which are taken to the colony, cut into tiny pieces and placed in fungal gardens. Workers specialise in tasks according to their sizes. The largest ants cut stalks, smaller workers chew the leaves and the smallest tend the fungus. Leafcutter ants are sensitive enough to recognise the reaction of the fungus to different plant material, apparently detecting chemical signals from the fungus. If a particular type of leaf is toxic to the fungus the colony will no longer collect it. The ants feed on structures produced by the fungi called gongylidia. Symbiotic bacteria on the exterior surface of the ants produce antibiotics that kill bacteria that may harm the fungi.
Foraging ants travel distances of up to from their nest and usually find their way back using scent trails. Some ants forage at night. Day foraging ants in hot and arid regions face death by desiccation, so the ability to find the shortest route back to the nest reduces that risk. Diurnal desert ants (Cataglyphis fortis) use visual landmarks in combination with other cues to navigate. In the absence of visual landmarks, the closely related Sahara desert ant (Cataglyphis bicolor) navigates by keeping track of direction as well as distance travelled, like an internal pedometer that counts how many steps they take in each direction. They integrate this information to find the shortest route back to their nest. Several species of ants are able to use the Earth's magnetic field. Ants' compound eyes have specialised cells that detect polarised light from the Sun, which is used to determine direction. These polarization detectors are sensitive in the ultraviolet region of the light spectrum.
Worker ants do not have wings and reproductive females lose their wings after their mating flights in order to begin their colonies. Therefore, unlike their wasp ancestors, most ants travel by walking. Some species are capable of leaping. For example, Jerdon's jumping ant (Harpegnathos saltator) is able to jump by synchronising the action of its mid and hind pairs of legs. There are several species of gliding ant including Cephalotes atratus; this may be a common trait among most arboreal ants. Ants with this ability are able to control the direction of their descent while falling.
Other species of ants can form chains to bridge gaps over water, underground, or through spaces in vegetation. Some species also form floating rafts that help them survive floods. These rafts may also have a role in allowing ants to colonise islands. Polyrhachis sokolova, a species of ant found in Australian mangrove swamps, can swim and live in underwater nests. Since they lack gills, they breathe in trapped pockets of air in the submerged nests.
Meat-eater ants feeding on a cicada. Social ants cooperate and collectively gather food.
Not all ants have the same kind of societies. The Australian bulldog ants are among the biggest and most basal (primitive) of ants. Like virtually all ants they are eusocial, but their social behaviour is poorly developed compared to other species. Each individual hunts alone, using its large eyes instead of its chemical senses to find prey.
Some species (such as Tetramorium caespitum) attack and take over neighbouring ant colonies. Others are less expansionist but just as aggressive; they invade colonies to steal eggs or larvae, which they either eat or raise as workers/slaves. Extreme specialists among these slave-raiding ants, such as the Amazon ants, are incapable of feeding themselves and need captured workers to survive. Captured workers of the enslaved species Temnothorax have evolved a counter strategy, destroying just the female pupae of the slave-making Protomognathus americanus, but sparing the males (who don't take part in slave-raiding as adults). See also New Scientist, 2009 April 9
A worker Harpegnathos saltator (a jumping ant) engaged in battle with a rival colony's queen.
Ants identify kin and nestmates through their scent, which comes from hydrocarbon-laced secretions that coat their exoskeletons. If an ant is separated from its original colony, it will eventually lose the colony scent. Any ant that enters a colony without a matching scent will be attacked.
Parasitic ant species enter the colonies of host ants and establish themselves as social parasites; species like Strumigenys xenos are entirely parasitic and do not have workers, but instead rely on the food gathered by their Strumigenys perplexa hosts. This form of parasitism is seen across many ant genera, but the parasitic ant is usually a species that is closely related to its host. A variety of methods are employed to enter the nest of the host ant. A parasitic queen can enter the host nest before the first brood has hatched, establishing herself prior to development of a colony scent. Other species use pheromones to confuse the host ants or to trick them into carrying the parasitic queen into the nest. Some simply fight their way into the nest. Hölldobler & Wilson (1990), pp. 436 448
A conflict between the sexes of a species is seen in some species of ants with the reproductives apparently competing to produce offspring that are as closely related to them as possible. The most extreme form involves the production of clonal offspring. An extreme of sexual conflict is seen in Wasmannia auropunctata, where the queens produce diploid daughters by thelytokous parthenogenesis and males produce clones by a process where a diploid egg loses its maternal contribution to produce haploid males that are clones of the father.
The spider Myrmarachne plataleoides (here a female) mimics weaver ants to avoid predators.
Army ants are nomadic and notorious for "raids", in which huge numbers of ants forage simultaneously over a certain area, attacking prey en masse. " The Savage, Beautiful World of Army Ants". National Public Radio (NPR). Armies of as many as 1,500,000 such ants destroy almost all animal life they encounter. Hymenopteran (insect). Encyclopædia Britannica. Dorylus sp., known as Siafu, " Driver ant, locally known as 'Siafu'". BBC - Science & Nature. attack everything in their path, including human beings. " Driver ant (insect)". Encyclopædia Britannica. Eciton burchellii is the swarming ant most commonly attended by "ant-following" birds such as antbirds and woodcreepers. Willis, E. & Y. Oniki (1978). "Birds and Army Ants" Annual Review of Ecology and Systematics 9: 243-263 Abstract
Ants form symbiotic associations with a range of species, including other ant species, other insects, plants, and fungi. They are preyed on by many animals and even certain fungi. Some arthropod species spend part of their lives within ant nests, either preying on ants, their larvae and eggs, consuming the ants' food stores, or avoiding predators. These inquilines can bear a close resemblance to ants. The nature of this ant mimicry (myrmecomorphy) varies, with some cases involving Batesian mimicry, where the mimic reduces the risk of predation. Others show Wasmannian mimicry, a form of mimicry seen only in inquilines.
An ant collects honeydew from an aphid.
Aphids and other hemipteran insects secrete a sweet liquid called honeydew when they feed on plant sap. The sugars in honeydew are a high-energy food source, which many ant species collect. In some cases the aphids secrete the honeydew in response to the ants' tapping them with their antennae. The ants in turn keep predators away and will move the aphids between feeding locations. On migrating to a new area, many colonies will take the aphids with them, to ensure a continued supply of honeydew. Ants also tend mealybugs to harvest their honeydew. Mealybugs can become a serious pest of pineapples if ants are present to protect mealybugs from their natural enemies.
Myrmecophilous (ant-loving) caterpillars of the family Lycaenidae (e.g., blues, coppers, or hairstreaks) are herded by the ants, led to feeding areas in the daytime, and brought inside the ants' nest at night. The caterpillars have a gland which secretes honeydew when the ants massage them. Some caterpillars produce vibrations and sounds that are perceived by the ants. Other caterpillars have evolved from ant-loving to ant-eating: these myrmecophagous caterpillars secrete a pheromone that makes the ants act as if the caterpillar is one of their own larvae. The caterpillar is then taken into the ants' nest where it feeds on the ant larvae.
Fungus-growing ants that make up the tribe Attini, including leafcutter ants, cultivate certain species of fungus in the Leucoagaricus or Leucocoprinus genera of the Agaricaceae family. In this ant-fungus mutualism, both species depend on each other for survival. The ant Allomerus decemarticulatus has evolved a three-way association with the host plant Hirtella physophora (Chrysobalanaceae), and a sticky fungus which is used to trap their insect prey.
Lemon ants make devil's gardens by killing surrounding plants with their stings and leaving a pure patch of lemon ant trees (Duroia hirsuta). This modification of the forest provides the ants with more nesting sites inside the stems of the Duroia trees. Some trees have extrafloral nectaries that provide food for ants, which in turn protect the plant from herbivorous insects. Species like the bullhorn acacia (Acacia cornigera) in Central America have hollow thorns that house colonies of stinging ants (Pseudomyrmex ferruginea) that defend the tree against insects, browsing mammals, and epiphytic vines. Isotopic labelling studies suggest that plants also obtain nitrogen from the ants. In return, the ants obtain food from protein and lipid rich Beltian bodies. Another example of this type of ectosymbiosis comes from the Macaranga tree, which has stems adapted to house colonies of Crematogaster ants.
Many tropical tree species have seeds that are dispersed by ants. Seed dispersal by ants or myrmecochory is widespread particularly in Africa and Australia. Some plants in fire-prone grassland systems are particularly dependent on ants for their survival and dispersal. Many ant-dispersed seeds have special external structures, elaiosomes, that are sought after by ants as food. A convergence, possibly a form of mimicry, is seen in the eggs of stick insects. They have an edible elaiosome-like structure and are taken into the ant nest where the young hatch.
A Meat ant tending a common leafhopper nymph
Ants prey on and obtain food from a number of social insects including other ants. Some species specialise in preying on termites (Megaponera and Termitopone) while a few Cerapachyinae prey on other ants. Some termites, including Nasutitermes corniger, form associations with certain ant species to keep away other predatory ant species. The tropical wasp Mischocyttarus drewseni coats the pedicel of its nest with an ant-repellant chemical. It is suggested that many tropical wasps may build their nests in trees and cover them to protect themselves from ants. Stingless bees (Trigona and Melipona) use chemical defences against ants.
Flies in the Old World genus Bengalia (Calliphoridae) prey on ants and are kleptoparasites, snatching prey or brood from the mandibles of adult ants. Wingless and legless females of the Malaysian phorid fly (Vestigipoda myrmolarvoidea) live in the nests of ants of the genus Aenictus and are cared for by the ants.
Fungi in the genera Cordyceps and Ophiocordyceps infect ants, causing them to climb up plants and sink their mandibles into plant tissue. The fungus kills the ant, grows on its remains, and produces a fruiting body. It appears that the fungus alters the behaviour of the ant to help disperse its spores in a microhabitat that best suits the fungus. Strepsipteran parasites also manipulate their ant host to climb grass stems, to help the parasite find mates. A nematode (Myrmeconema neotropicum) that infects canopy ants (Cephalotes atratus) causes the black coloured gasters of workers to turn red. The parasite also alters the behaviour of the ant, and makes them carry their gasters high. The conspicuous red gasters are mistaken by birds for ripe fruits such as Hyeronima alchorneoides and eaten. The droppings of the bird are collected by other ants and fed to their young leading to the further spread of the nematode.
Spiders also feed on ants
South American poison dart frogs in the genus Dendrobates feed mainly on ants, and the toxins in their skin may come from the ants. Several South American antbirds follow army ants to feed on the insects that are flushed from cover by the foraging ants.
This behaviour was once considered mutualistic, but later studies show that it is instead kleptoparasitic, with the birds stealing prey.
Birds indulge in a peculiar behaviour called anting that is as yet not fully understood. Here birds rest on ant nests, or pick and drop ants onto their wings and feathers; this may remove ectoparasites. Anteaters, pangolins and several marsupial species in Australia have special adaptations for living on a diet of ants. These adaptations include long, sticky tongues to capture ants and strong claws to break into ant nests. Brown bears (Ursus arctos) have been found to feed on ants, and about 12%, 16%, and 4% of their faecal volume in spring, summer, and autumn, respectively, is composed of ants.
Weaver ants are used as a biological control for citrus cultivation in southern China.
Ants perform many ecological roles that are beneficial to humans, including the suppression of pest populations and aeration of the soil. The use of weaver ants in citrus cultivation in southern China is considered one of the oldest known applications of biological control. Hölldobler & Wilson (1990), pp. 619 629 On the other hand, ants can become nuisances when they invade buildings, or cause economic losses.
In some parts of the world (mainly Africa and South America), large ants, especially army ants, are used as surgical sutures. The wound is pressed together and ants are applied along it. The ant seizes the edges of the wound in its mandibles and locks in place. The body is then cut off and the head and mandibles remain in place to close the wound.
Some ants of the family Ponerinae have toxic venom and are of medical importance. The species include Paraponera clavata (Tocandira) and Dinoponera spp. (false Tocandiras) of South America and the Myrmecia ants of Australia.
In South Africa, ants are used to help harvest rooibos (Aspalathus linearis), which are small seeds used to make a herbal tea. The plant disperses its seeds widely, making manual collection difficult. Black ants collect and store these and other seeds in their nest, where humans can gather them en masse. Up to half a pound (200 g) of seeds can be collected from one ant-heap.
Although most ants survive attempts by humans to eradicate them, a few are highly endangered. These are mainly island species that have evolved specialized traits and include the critically endangered Sri Lankan relict ant (Aneuretus simoni) and Adetomyrma venatrix of Madagascar.
Ant larvae on sale in Isaan, Thailand
Ants and their larvae are eaten in different parts of the world. The eggs of two species of ants are the basis for the dish in Mexico known as escamoles. They are considered a form of insect caviar and can sell for as much as USD 40 per pound (USD 90/kg) because they are seasonal and hard to find. In the Colombian department of Santander, hormigas culonas (roughly interpreted as "large-bottomed ants") Atta laevigata are toasted alive and eaten.
In areas of India, and throughout Burma and Thailand, a paste of the green weaver ant (Oecophylla smaragdina) is served as a condiment with curry. Weaver ant eggs and larvae as well as the ants themselves may be used in a Thai salad, yum (ยำ), in a dish called yum khai mod daeng (ยำà¹à¸à¹à¸¡à¸à¹à¸à¸) or red ant egg salad, a dish that comes from the Issan or north-eastern region of Thailand. Saville-Kent, in the Naturalist in Australia wrote "Beauty, in the case of the green ant, is more than skin-deep. Their attractive, almost sweetmeat-like translucency possibly invited the first essays at their consumption by the human species". Mashed up in water, after the manner of lemon squash, "these ants form a pleasant acid drink which is held in high favor by the natives of North Queensland, and is even appreciated by many European palates".
In his First Summer in the Sierra, John Muir notes that the Digger Indians of California ate the tickly acid gasters of the large jet-black carpenter ants. The Mexican Indians eat the replete workers, or living honey-pots, of the honey ant (Myrmecocystus).
The tiny pharaoh ant is a major pest in hospitals and office blocks; it can make nests between sheets of paper.
Some ant species are considered pests, and because of the adaptive nature of ant colonies, eliminating the entire colony is nearly impossible. Pest management is therefore a matter of controlling local populations, instead of eliminating an entire colony, and most attempts at control are temporary solutions.
Ants classified as pests include the pavement ant, yellow crazy ant, sugar ants, the Pharaoh ant, carpenter ants, Argentine ant, odorous house ants, red imported fire ant and European fire ant. Populations are controlled using insecticide baits, either in granule or liquid formulations. Bait is gathered by the ants as food and brought back to the nest where the poison is inadvertently spread to other colony members through trophallaxis. Boric acid and borax are often used as insecticides that are relatively safe for humans. Bait may be broadcast over a large area to control species like the red fire ant that occupy large areas. Nests of red fire ants may be destroyed by following the ants' trails back to the nest and then pouring boiling water into it to kill the queen. This works in about 60% of the mounds and requires about per mound.
Myrmecologists study ants in the laboratory and in their natural conditions. Their complex and variable social structures have made ants ideal model organisms. Ultraviolet vision was first discovered in ants by Sir John Lubbok in 1881. Studies on ants have tested hypotheses in ecology, sociobiology and have been particularly important in examining the predictions of theories of kin selection and evolutionarily stable strategies. Ant colonies can be studied by rearing or temporarily maintaining them in formicaria, specially constructed glass framed enclosures. Individuals may be tracked for study by marking them with colours.
The successful techniques used by ant colonies have been studied in computer science and robotics to produce distributed and fault-tolerant systems for solving problems. This area of biomimetics has led to studies of ant locomotion, search engines that make use of "foraging trails", fault-tolerant storage and networking algorithms.
Aesop's ants: picture by Milo Winter, 1888 1956
Ants have often been used in fables and children's stories to represent industriousness and cooperative effort. They are also mentioned in religious texts. In the Book of Proverbs in the Bible, ants are held up as a good example for humans for their hard work and cooperation. Aesop did the same in his fable The Ant and the Grasshopper. In the Quran, Sulayman ( ) is said to have heard and understood an ant warning other ants to return home to avoid being accidentally crushed by Sulayman and his marching army. In parts of Africa, ants are considered to be the messengers of the gods. Ant bites are often said to have curative properties. The sting of some species of Pseudomyrmex is claimed to give fever relief. Some Native American mythology, such as the Hopi mythology, considers ants as the very first animals. Others use ant bites in initiation ceremonies as a test of endurance.
Ant society has always fascinated humans and has been written about both humorously and seriously. Mark Twain wrote about ants in his A Tramp Abroad. Some modern authors have used the example of the ants to comment on the relationship between society and the individual. Examples are Robert Frost in his poem "Departmental" and T. H. White in his fantasy novel The Once and Future King. The plot in French entomologist and writer Bernard Werber's Les Fourmis science-fiction trilogy is divided between the worlds of ants and humans; ants and their behaviour is described using contemporary scientific knowledge. In more recent times, animated cartoons and 3D animated movies featuring ants have been produced include Antz, A Bug's Life, The Ant Bully, The Ant and the Aardvark, Atom Ant, and there is a comic book superhero called Ant-Man.
The Chinese character for ant (è»/è) is a combination of logograms that may be interpreted as "insect (è«) which behaves properly (義/ä¹)". The Japanese character for ant (è») also shares this etymology.
From the late 1950s through the late 1970s, ant farms were popular educational children's toys in the United States. Later versions use transparent gel instead of soil allowing greater visibility. In the early 1990s, the video game SimAnt, which simulated an ant colony, won the 1992 Codie award for "Best Simulation Program".
Ants are also quite popular inspiration for many science-fiction creatures, such as the Formics of Ender's Game, the Bugs of Starship Troopers, the giant ants in the film Them!, and ants mutated into super intelligence in Phase IV. In strategy games, ant-based species often benefit from increased production rates due to their single-minded focus, such as the Klackons in the Master of Orion series of games or the ChCht in Deadlock II. These characters are often credited with a hive mind, a common misconception about ant colonies.
* Ant robotics
* Ant stings
* International Union for the Study of Social Insects
* Myrmecological News (journal)
* Task allocation and partitioning of social insects
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* Antweb from The California Academy of Sciences
* AntBlog a website dedicated to the study of ant colonies
* AntBase - a taxonomic database with literature sources
* Discover Life images, information and links
* BugGuide
* Global Ant Project
* Navajo Ant Project
* Gakushu Kenkyusha (1979) Ants. Gakken's Photo Encyclopedia
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Antwerp | Is Antwerp a city? | Yes | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Is Antwerp a city? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Is Antwerp a municipality? | Yes | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Is Antwerp a municipality? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Is Antwerp in Belgium? | Yes | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Is Antwerp in Belgium? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | What is Antwerp? | Antwerp is a city and municipality in Belgium. | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | What is Antwerp? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | What is the population of the city of Antwerp? | Antwerp's population is 472,071. | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | What is the population of the city of Antwerp? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Where is the city of Antwerp? | Antwerp is in Belgium | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Where is the city of Antwerp? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Why is Antwerp important to fashion? | Yes | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Why is Antwerp important to fashion? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Antwerp is to the east of what river? | Antwerp is to the east of the Scheldt river | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | Antwerp is to the east of what river? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | How many municipalities does Antwerp have? | None, Antwerp is a municipality. | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Antwerp | How many municipalities does Antwerp have? | null | data/set3/a4 | Antwerp
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|Foreground: Statue of the giant's hand being thrown into the Scheldt River. Background: Town hall
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady) and the Scheldt river.
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|Grote Markt
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Antwerp ( , Dutch: , ) is a city and municipality in Belgium and the capital of the Antwerp province in Flanders, one of Belgium's three regions. Antwerp's total population is 472,071 (as of 1 January 2008) Statistics Belgium; Population de droit par commune au 1 janvier 2008 (excel-file) Population of all municipalities in Belgium, as of 1 January 2008. Retrieved on 2008-10-19. and its total area is , giving a population density of 2,308 inhabitants per km². The metropolitan area, including the outer commuter zone, covers an area of with a total of 1,190,769 inhabitants as of 1 January 2008. Statistics Belgium; De Belgische Stadsgewesten 2001 (pdf-file) Definitions of metropolitan areas in Belgium. The metropolitan area of Antwerp is divided into three levels. First, the central agglomeration (agglomeratie) with 715,301 inhabitants (2008-01-01). Adding the closest surroundings (banlieue) gives a total of 955,338. And, including the outer commuter zone (forensenwoonzone) the population is 1,190,769. Retrieved on 2008-10-19.
Antwerp has long been an important city in the nations of the Benelux both economically and culturally, especially before the Spanish Fury of the Dutch Revolt. It is located on the right bank of the river Scheldt, which is linked to the North Sea by the estuary Westerschelde.
According to folklore, and as celebrated by the statue in front of the town hall, the city got its name from a legend involving a mythical giant called Antigoon who lived near the river Scheldt. He exacted a toll from those crossing the river, and for those who refused, he severed one of their hands and threw it into the river Scheldt. Eventually, the giant was slain by a young hero named Brabo, who cut off the giant's own hand and flung it into the river. Hence the name Antwerpen, from Dutch hand werpenâakin to Old English hand and wearpan (= to throw), that has changed to today's warp. Brabo Antwerpen 1 (centrum) / Antwerpen
In favour of this folk etymology is the fact that hand-cutting was indeed practised in Europe, the right hand of a man who died without issue being cut off and sent to the feudal lord as proof of main-morte.
However, John Lothrop Motley argues that Antwerp's name derives from an 't werf (on the wharf).
Aan 't werp (at the warp) is also possible. This 'warp' (thrown ground) would be a man made hill, just high enough to remain dry at high tide, whereupon a farm would be built. Another word for werp is pol (hence polders).
The most prevailing theory is that the name originated in the Gallo-Roman period and comes from the Latin antverpia. Antverpia would come from Ante (before) Verpia (deposition, sedimentation), indicating land that forms by deposition in the inside curve of a river. Note that the river Scheldt, before a transition period between 600 to 750, followed a different track. This must have coincided roughly with the current ringway south of the city, situating the city within a former curve of the river. Antwerp Tourist Information - Meredith Booney, "The name 'Antwerp' has been linked to the word "aanwerp" (alluvial mound), which was the geographical feature in the early settlement period in this place".
Historical Antwerp had its origins in a Gallo-Roman vicus civilization. Excavations carried out in the oldest section near the Scheldt, 1952-1961 (ref. Princeton), produced pottery shards and fragments of glass from mid-second century to the end of the third century.
In the 4th century, Antwerp was first named, having been settled by the Germanic Franks. Brittanica: Antwerp The name was reputed to have been derived from "anda" (at) and "werpum" (wharf).
The Merovingian Antwerp, now fortified, was evangelized by Saint Amand in the 7th century. At the end of the 10th century, the Scheldt became the boundary of the Holy Roman Empire. Antwerp became a margraviate, a border province facing the County of Flanders.
In the 11th century Godfrey of Bouillon was for some years known as the marquis of Antwerp. In the 12th century, Norbert of Xanten established a community of his Premonstratensian canons at St. Michael's Abbey at Caloes. Antwerp was also the headquarters of Edward III during his early negotiations with Jacob van Artevelde, and his son Lionel, the earl of Cambridge, was born there in 1338.
After the closing of the Zwin and the consequent decline of Bruges, the city of Antwerp, then part of the Duchy of Brabant, became of importance. At the end of the 15th century the foreign trading houses were transferred from Bruges to Antwerp, and the building assigned to the English nation is specifically mentioned in 1510.
Fernand Braudel states that Antwerp became "the center of the entire international economy, something Bruges had never been even at its height." (Braudel 1985 p. 143.) Antwerp was the richest city in Europe at this time. Antwerp's golden age is tightly linked to the "Age of Exploration". Over the first half of the 16th century Antwerp grew to become the second-largest European city north of the Alps by 1560. Many foreign merchants were resident in the city. Francesco Guicciardini, the Venetian envoy, stated that hundreds of ships would pass in a day, and 2,000 carts entered the city each week. Portuguese ships laden with pepper and cinnamon would unload their cargo.
Without a long-distance merchant fleet, and governed by an oligarchy of banker-aristocrats forbidden to engage in trade, the economy of Antwerp was foreigner-controlled, which made the city very cosmopolitan, with merchants and traders from Venice, Ragusa, Spain and Portugal. Antwerp had a policy of toleration, which attracted a large orthodox Jewish community. Antwerp was not a "free" city though, since it had been reabsorbed into the Duchy of Brabant in 1406 and was controlled from Brussels.
Antwerp experienced three booms during its golden age: The first based on the pepper market, a second launched by American silver coming from Seville (ending with the bankruptcy of Spain in 1557), and a third boom, after the stabilising Treaty of Cateau-Cambresis, in 1559, based on the textiles industry. The boom-and-bust cycles and inflationary cost-of-living squeezed less-skilled workers.
The religious revolution of the Reformation erupted in violent riots in August 1566, as in other parts of the Netherlands. The regent Margaret, Duchess of Parma, was swept aside when Philip II sent the Duke of Alba at the head of an army the following summer. When the Eighty Years' War broke out in 1572, commercial trading between Antwerp and the Spanish port of Bilbao collapsed and became impossible. On November 4, 1576, Spanish soldiers plundered the city. During the Spanish Fury 6,000 citizens were massacred, 800 houses were burnt down, and over 2 million sterling of damage was done.
Antwerp became the capital of the Dutch revolt. In 1585, Alessandro Farnese, Duke of Parma and Piacenza, captured it after a long siege and as part of the terms of surrender its Protestant citizens were given two years to settle their affairs before quitting the city. Boxer Charles Ralph, The Dutch seaborne empire, 1600-1800, p. 18, Taylor & Francis, 1977 ISBN 0091310512, 9780091310516
Google books Most went to the United Provinces in the north starting the Dutch Golden Age. Antwerp's banking was controlled for a generation by Genoa, and Amsterdam became the new trading centre.
Map of Antwerp, its buildings and the march. (1624)
The recognition of the independence of the United Provinces by the Treaty of Münster in 1648 stipulated that the Scheldt should be closed to navigation, which destroyed Antwerp's trading activities. This impediment remained in force until 1863, although the provisions were relaxed during French rule from 1795 to 1814, and also during the time Belgium formed part of the Kingdom of the United Netherlands (1815 to 1830). Antwerp had reached the lowest point of its fortunes in 1800, and its population had sunk under 40,000, when Napoleon, realizing its strategic importance, assigned two million to enlarge the harbor by constructing two docks and a mole and deepening the Scheldt to allow for larger ships to approach Antwerp. Napoleon hoped that by making Antwerp's harbor the finest in Europe he would be able to counter London's harbor and stint English growth, but he was defeated at the Battle of Waterloo before he could see the plan through.
In 1830, the city was captured by the Belgian insurgents, but the citadel continued to be held by a Dutch garrison under General David Hendrik Chassé. For a time Chassé subjected the town to periodic bombardment which inflicted much damage, and at the end of 1832 the citadel itself was besieged by a French army. During this attack the town was further damaged. In December 1832, after a gallant defence, Chassé made an honourable surrender.
Antwerp was the first city to host the World Gymnastics Championships, in 1903. During World War I, the city became the fallback point of the Belgian Army after the defeat at Liège. It was taken after heavy fighting by the German Army, and the Belgians were forced to retreat westward.
Antwerp hosted the 1920 Summer Olympics. During World War II, the city was an important strategic target because of its port. It was occupied by Germany in May 1940 and liberated by the British 11th Armoured Division on September 4, 1944. After this, the Germans attempted to destroy the Port of Antwerp, which was used by the Allies to bring new material ashore. Thousands of V-1 and V-2 missiles battered the city. The city was hit by more V-2s than all other targets during the entire war combined, but the attack did not succeed in destroying the port since many of the missiles fell upon other parts of the city. As a result, the city itself was severely damaged and rebuilt after the war in a modern style. After the war, Antwerp, which had already had a sizable Jewish population before the war, once again became a major European center of Haredi (and particularly Hasidic) Orthodox Judaism.
Population time-line of Antwerp.
This is the population of the city of Antwerp only, not of the larger current municipality of the same name.
Districts of Antwerp.
The municipality comprises the city of Antwerp proper and several towns. It is divided into nine entities (districts):
#Antwerp (district)
#Berchem
#Berendrecht-Zandvliet-Lillo
#Borgerhout
#Deurne
#Ekeren
#Hoboken
#Merksem
#Wilrijk
|Antwerp City Hall at the Grote Markt (Main Square).
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|16th-century Guildhouses at the Grote Markt.
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|The Onze-Lieve-Vrouwekathedraal (Cathedral of our Lady), here seen from the Groenplaats, is the highest cathedral in the Low Countries and home to several triptychs by Baroque painter Rubens. It remains the tallest building in the city.
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|Statue of Brabo and the giant's hand
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|Antwerp lawcourts
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In the 16th century, Antwerp was noted for the wealth of its citizens ("Antwerpia nummis"); the houses of these wealthy merchants and manufacturers have been preserved throughout the city. However fire has destroyed several old buildings, such as the house of the Hanseatic League on the northern quays in 1891. The city also suffered considerable war damage by V-bombs, and in recent years other noteworthy buildings were demolished for new developments.
*Antwerp Zoo was founded in 1843, and is home to more than 6,000 animals (about 769 species). One of the oldest zoos in the world, it is renowned for of its high level of research and conservation.
*Central Station is a railway station designed by Louis Delacenserie that was completed in 1905. It has two monumental neo-baroque facades, a large metal and glass dome (60m/197 ft) and a gilt and marble interior
*Cathedral of Our Lady. This church was begun in the 14th century and finished in 1518. The church has four works by Rubens, viz. "The Descent from the Cross", "The Elevation of the Cross", "The Resurrection of Christ" and "The Assumption"
*Saint James' Church, is more ornate than the cathedral. It contains the tomb of Rubens
*Church of St Paul, has a beautiful baroque interior. It is a few hundred yards north of the Grote Markt
* Plantin-Moretus Museum preserves the house of the printer Christoffel Plantijn and his successor Jan Moretus
*Boerentoren (Farmers' Tower) or KBC Tower, a 26-storey building built in 1932, is the oldest skyscraper in Europe Emporis. Retrieved October 23, 2006.
*Royal Museum of Fine Arts, close to the southern quays, has a collection of old masters (Rubens, Van Dyck, Titian) and the leading Dutch masters.
* Rubenshuis is the former home and studio of Peter Paul Rubens (1577-1640) in Antwerp. It is now a museum.
* exchange or Bourse, one of the earliest institutions in Europe with that title, was built in 1872.
*law courts, designed by the Richard Rogers Partnership, Arup and VK Studio, and opened by King Albert in April 2006. This building is the antithesis of the heavy, dark court building designed by Joseph Poelaert that dominates the skyline of Brussels. The courtrooms sit on top of six fingers that radiate from an airy central hall, and are surmounted by spires which provide north light and resemble oast houses or the sails of barges on the nearby River Scheldt. It is built on the site of the old Zuid ("South") station, at the end of a magnificent 1.5 km perspective at the southern end of Amerikalei. The road neatly disappears into an underpass under oval Bolivarplaats to join the motorway ring. This leaves peaceful surface access by foot, bicycle or tram (routes 8 & 12). The building's highest 'sail' is 51 m high, has a floor area of 77,000 m², and cost â¬130m.
Het Steen (literally: 'The Stone').
Although Antwerp was formerly a fortified city, nothing remains of the former enceinte or of the old citadel defended by General Chassé in 1832, except for the Steen, which has been restored. Modern Antwerp's broad avenues mark the position of the original fortifications. After the establishment of Belgian independence, Antwerp was defended by the citadel and an enceinte around the city. In 1859, seventeen of the twenty-two fortresses constructed under Wellington's supervision in 1815â1818 were dismantled and the old citadel and enceinte were removed. A new enceinte long was constructed, and the villages of Berchem and Borgerhout, now boroughs of Antwerp, were absorbed within the city.
This enceinte is protected by a broad wet ditch, and in the caponiers are the magazines and store chambers of the fortress. The enceinte has nineteen openings or gateways, but of these seven are not used by the public. As soon as the enceinte was finished eight detached forts from 2 to 2-½ miles from the enceinte were constructed. They begin on the north near Wijnegem and the zone of inundation, and terminate on the south at Hoboken. In 1870 Fort Merksem and the redoubts of Berendrecht and Oorderen were built for the defence of the area to be inundated north of Antwerp.
In the 1870s, the fortifications of Antwerp were deemed to be out of date, given the increased range and power of artillery and explosives. Antwerp was transformed into a fortified position by constructing an outer line of forts and batteries 6 to from the enceinte.
The Boerentoren ("Farmers' tower"), nickname of the KBC Bank building in Antwerp.
According to the American Association of Port Authorities (AAPA), the port of Antwerp was the seventeenth largest (by tonnage) port in the world in 2005 and second only to Rotterdam in Europe. Importantly it handles high volumes of economically attractive general and project cargo, as well as bulk cargo. Antwerp's docklands, with five oil refineries, are home to a massive concentration of petrochemical industries, second only to the petrochemical cluster in Houston, Texas. Electricity generation is also an important activity, with four nuclear power plants at Doel, a conventional power station in Kallo, as well as several smaller combined cycle plants. There are plans for a wind farm in a disused area of the docklands.
The old Belgian bluestone quays bordering the Scheldt for a distance of to the north and south of the city centre have been retained for their sentimental value and are used mainly by cruise ships and short-sea shipping.
Antwerp's other great mainstay is the diamond trade. The city has four diamond bourses: one for bort and three for gem quality goods. Since World War II families of the large Hasidic Jewish community have dominated Antwerp's diamond trading industry, although the last two decades have seen Indian and Armenian traders become increasingly important.
Antwerp World Diamond Centre, the successor to the Hoge Raad voor Diamant, plays an important role in setting standards, regulating professional ethics, training and promoting the interests of Antwerp as a centre of the diamond industry.
A motorway bypass encircles much of the city centre. Known locally as the "Ring" it offers motorway connections to Brussels, Hasselt and Liège, Ghent, Lille and Bruges and Breda and Bergen op Zoom (Netherlands). The banks of the Scheldt are linked by three road tunnels (in order of construction): the Waasland Tunnel (1934), the Kennedy Tunnel (1967) and the Liefkenshoek Tunnel (1991). Currently a fourth high volume highway link called "Oosterweelconnection" is in the tendering stage. It will entail the construction of a long viaduct and bridge (the Lange Wapper Bridge) over the Scheldt on the north side of the city. The completion date is as yet uncertain. The cost of the connection is estimated at 2.2 billion euro.
Antwerp is the focus of lines to the north to Essen and the Netherlands, east to Turnhout, south to Mechelen, Brussels and Charleroi via Luttre, and southwest to Ghent and Ostend. It is served by international trains to Amsterdam and Paris, and national trains to Ghent, Bruges, Ostend, Brussels, Charleroi, Hasselt, Liège and Turnhout.
Antwerp's Central station is an architectural monument in itself, and is mentioned in W G Sebald's haunting novel Austerlitz. Prior to the completion in 2007 of a tunnel that runs northwards under the city centre to emerge at the old Antwerp Dam station, Centraal was a terminus. Trains to the Netherlands either had to reverse at Centraal or call only at Berchem station, 2 km to the south, and then describe a semicircle to the east, round the Singel.
The city has a web of tram and bus lines operated by De Lijn and providing access to the city centre, suburbs and the Left Bank. The tram network has 12 lines, of which the underground section is called the "premetro" and includes a tunnel under the river.
Antwerp International Airport is in the district of Deurne. VLM Airlines flies to London (City Airport) and Manchester in England and remains the only airline with scheduled air services to and from Antwerp International Airport. The airport is connected by bus to the city center. Brussels Airport is about 45 km from the city of Antwerp, and connects the city worldwide. The airport is connected by bus and by train to the city centre of Antwerp.
One of the many Marian statues which feature on Antwerp street corners
Antwerp had an artistic reputation in the 17th century, based on its school of painting, which included Rubens, Van Dyck, Jordaens, the two Teniers and many others. Informally, most Antverpians (in Dutch Antwerpenaren, people from Antwerp) daily speak Antverpian (in Dutch Antwerps), a dialect that Dutch-speakers know as distinctive from other Brabantic dialects through its typical vowel pronunciations: approximating the vowel sound in 'bore'â for one of its long 'a'-sounds while other short 'a's are very sharp like the vowel sound in 'hat'. The Echt Antwaarps Teater ("Authentic Antverpian Theatre") brings the dialect on stage.
Antwerp is a rising fashion city, and has produced designers such as the Antwerp Six. The city has a cult status in the fashion world, due to the Royal Academy of Fine Arts, one of the most important fashion academies in Europe. It has served as the learning centre for a large number of Belgian fashion designers. Since the 1980s, several graduates of the Belgian Royal Academy of Fine Arts have become internationally successful fashion designers in Antwerp.
Antwerp is famous for its local products and in August every year the Bollekesfeest takes place. The Bollekesfeest is a showcase for such local products as beer from the De Koninck Brewery, better known in Antwerp as a "Bolleke", the Mokatine sweets made by Confiserie Roodthooft, Elixir D'Anvers, a locally-made liqueur, locally roasted coffee from Koffie Verheyen, sugar from Candico, Poolster pickled herring, Equinox horse meat, and others. The local products are represented by a non-profit making organisation, Streekproducten Provincie Antwerpen vzw.
The major sport clubs are K.F.C. Germinal Beerschot and R. Antwerp F.C. (football) and Antwerp Diamond Giants (basketball).
After the Holocaust and the destruction of its many semi-assimilated Jews, Antwerp became a major centre for Orthodox Jews. At present, about 15,000 Haredi Jews, mostly Hasidic, live in Antwerp. The city has three official Jewish Congregations: Shomrei Hadass, headed by Rabbi Dovid Moishe Lieberman, Machsike Hadass, headed by Rabbi Eliyahu Sternbuch (formerly Chief Rabbi Chaïm Kreiswirth) and the Portuguese Community Bne Moshe. Antwerp has an extensive network of synagogues, shops, schools and organizations, within the Machsike Hadas community. Significant Hasidic movements in Antwerp include Pshevorsk, based in Antwerp, as well as branches of Satmar, Belz, Bobov, Ger, Skver, Klausenburg and several others. Rabbi Chaim Kreiswirth, chief rabbi of the Machsike Hadas community, who died in 2003, was arguably one of the better known personalities to have been based in Antwerp. An attempt to have a street named after him has received the support of the Town Hall and is in the process of being implemented.
A number of Christian missions to seafarers are based in Antwerp, notably on the Italiëlei. These include the Mission to Seafarers, British & International Sailorsâ Society, the Finnish Seamen's Mission, the Norwegian Sjømannskirken and the Apostleship of the Sea. They provide cafeterias, cultural and social activities as well as religious services.
The following places are twinned with or sister cities to Antwerp:
Within the context of development cooperation, Antwerp is also linked to:
* Paramaribo, Suriname
* Durban, South Africa
Abraham Ortelius.
Hendrik Conscience
*Lionel of Antwerp, 1st Duke of Clarence, son of Edward III of England (1338â1368)
*Samuel Blommaert, Director of the Dutch West India Company (1583 â 1654)
*Frans Floris, painter (1520â1570)
*Abraham Ortelius, cartographer and geographer (1527â1598)
*Gillis van Coninxloo, painter of forest landscapes (1544â1607)
*Bartholomeus Spranger, painter, draughtsman, and etcher (1546â1611)
*Paul and Mattheus Brill, landscape painters (1554-1626, 1550-1583, resp.)
*Abraham Janssens, painter (c. 1570-1632)
*Rodrigo Calderón, Count of Oliva, Spanish favourite and adventurer (d. 1621)
*Frans Snyders, still life and animal painter (1579â1657)
*Frans Hals, painter (1580â1666)
*Caspar de Crayer, painter (1582â1669)
*David Teniers the Elder, painter (1582â1649)
*Jacob Jordaens, painter (1593â1678)
*Anthony van Dyck, painter (1599â1641)
*David Teniers the Younger, painter (1610â1690)
*Jan Fyt, animal painter (1611â1661)
*Nicolaes Maes, Baroque painter (1634â1693)
*Gerard Edelinck, copper-plate engraver (1649â1707)
*Peter Tillemans, painter (c. 1684â1734)
*John Michael Rysbrack, sculptor (1694â1770)
*Hendrik Conscience, writer and author of De Leeuw van Vlaanderen ("The Lion of Flanders") (1812â1883)
*Georges Eekhoud, novelist (1854â1927)
*Hippolyte Delehaye, Jesuit Priest and hagiographic scholar (1859â1941)
*Willem Elsschot, writer and poet (1882â1960)
*Constant Permeke, expressionist painter (1886â1952)
*Paul van Ostaijen, poet and writer (1896â1928)
*Albert Lilar, Minister of Justice (1900â1976)
*Maurice Gilliams, writer (1900â1982)
*Antoinette Feuerwerker, French jurist and member of the Resistance (1912-2003)
*Paul Buysse , businessman (1945 -)
*Evi Goffin, vocalist (1981- )
*Jessica Van Der Steen, Model (1984 -)
*Karl Gotch, professional wrestler (1924â2007)
*Tom Barman, Belgian musician and film director.
*Willem Usselincx , Flemish merchant and investor, one of the founders of the Dutch West India Company (1567-1647)
Joachim Patinir.
Wenceslas Hollar.
*Quentin Matsys, Renaissance painter, founder of the Antwerp school (1466â1530)
*Jan Mabuse, painter (c. 1478-1532)
*Joachim Patinir, landscape and religious painter (c. 1480-1524)
*John Rogers, minister of religion, Bible translator and commentator, and martyr (c. 1500-1555)
*Joos van Cleve, painter (c. 1500-1540/41)
*Damião de Góis, Portuguese humanist philosopher (1502â1574)
*Sir Thomas Gresham, English merchant and financier (c. 1519-1579)
*Sir Anthony More, portrait painter (1520- c. 1577)
*Christoffel Plantijn, humanist, book printer and publisher (c. 1520-1589)
*Pieter Brueghel the Elder, painter and printmaker (1525â1569)
*Philip van Marnix, writer and statesman (1538â1598)
*Simon Stevin, mathematician and engineer (c. 1548/49-1620)
*John Bull, English/Welsh composer, musician, and organ builder (c. 1562-1628)
*Jan Brueghel the Elder, also known as "Velvet" Brueghel, painter (1568â1625)
*Pieter Paul Rubens, painter (1577â1640)
*William Cavendish, 1st Duke of Newcastle, English soldier, politician, and writer (c. 1592-1676)
*Adriaen Brouwer, painter (1605â1638)
*Jan Davidszoon de Heem, painter (1606â1684)
*Wenceslas Hollar, Bohemian etcher (1607â1677)
*Jan Lievens, painter (1607â1674)
*Jan Frans Willems, writer (1793â1846)
*Henri Alexis Brialmont, military engineer (1821â1903)
*Sir Lawrence Alma-Tadema, painter (1836â1912)
*Vincent van Gogh, impressionist painter, lived in Antwerp for about four months (1853â1890)
*Camille Huysmans, Socialist politician and former Prime Minister of Belgium (1871â1968)
*Moshe Yitzchok Gewirtzman, leader of the Hasidic Pshevorsk movement based in Antwerp (1881â1976)
*Romi Goldmuntz, businessman (1882â1960)
*Gerard Walschap, writer (1898â1989)
*Albert Lilar, Minister of Justice (1900â1976)
*Suzanne Lilar, essayist, novelist, and playwright (1901â1992)
*Philip Sessarego, former British Army soldier, conman, hoaxer, mercenary lived in Antwerp and found dead in a garage (1952-2008)
*Jean Genet, French writer and political activist: lived in Antwerp for short period in 1930s (1909â1986)
*George du Maurier, Came to Antwerp to study art and lost the sight in one eye. Cartoonist, author and grandfather of Daphne du Maurier (1834â1896)
*Chaim Kreiswirth, Talmudist and Rabbi of the Machsike Hadas Community, Antwerp (1918â2001)
*William Tyndale, Bible translator, arrested in Antwerp 1535 and burnt at Vilvoorde in 1536 (ca. 1494-1536)
*Akiba Rubinstein, Polish grandmaster of chess (1882â1961).
*Veerle Casteleyn, Belgian performer
* Den Dam â an area in northern Antwerp
* Linkeroever - an area on the left bank of the Scheldt with a lot of apartment buildings
* Meir â Antwerp's largest shopping street
* Seefhoek - an area in north-east Antwerp, situated around the Stuyvenbergplein
* Van Wesenbekestraat â the Chinatown of Antwerp
* Zuid â the south of Antwerp
* Zurenborg
* Antwerp Book Fair
* Antwerp lace
* Antwerp Water Works (AWW)
* Archief en Museum voor het Vlaams Cultuurleven
* Jewish Community of Antwerp
* List of mayors of Antwerp
* Pshevorsk â Hassidic Jewish movement based in Antwerp
* University of Antwerp
* Carolus Scribani, Origines Antwerpiensium, 1610
* Gens, Histoire de la ville d'Anvers
* F.H. Mertens, K.L. Torfs, Geschiedenis van Antwerpen sedert de stichting der. stad tot onze tyden, vol. 7, Antwerp 1853
* J. L. Motley, Rise of the Dutch Republic, 1856
* P. Génard, Anvers à travers les ages
* Annuaire statistique de la Belgique
* Richard Stillwell, ed. Princeton Encyclopedia of Classical Sites, 1976: "Antwerp Belgium"
*
* Official Website
* Capsule History
* Tourism Antwerp
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Arabic_language | Is Arabic a Central Semitic language? | Yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Is Arabic a Central Semitic language? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Was Arabic a major vehicle of culture in Europe? | Yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Was Arabic a major vehicle of culture in Europe? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Is Arabic classified alongside Semitic languages? | Yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Is Arabic classified alongside Semitic languages? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | How many people speak the Arabic language? | 280 million people. | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | How many people speak the Arabic language? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | When was Arabic calligraphy invented? | Many styles were developed after 786. | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | When was Arabic calligraphy invented? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Where is Arabic spoken? | The Middle East and North Africa | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Where is Arabic spoken? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Why is Arabic related to Islam? | Arabic is the liturgical language of Islam | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Why is Arabic related to Islam? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Why does Arabic heavily influence European languages? | Arabic was a major vehicle of culture in Europe, and the Arab and European civilizations are geographically close. | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Why does Arabic heavily influence European languages? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | How do you say "black" in Arabic? | null | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Is Arabic the largest member of the Semitic language family? | yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Is Arabic the largest member of the Semitic language family? | yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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Arabic_language | Does Modern Standard Arabic continue to evolve like other languages? | yes | data/set5/a3 | Arabic_language
Arabic ( , ( ) or ) is a Central Semitic language, thus related to and classified alongside other Semitic languages such as Hebrew and the Neo-Aramaic languages. In terms of speakers, Arabic is the largest member of the Semitic language family. It is spoken by more than 280 million people as a first language, most of whom live in the Middle East and North Africa, and by 250 million more as a second language. Arabic has many different, geographically-distributed spoken varieties, some of which are mutually unintelligible. "Arabic language." Encyclopaedia Britannica. 2009. Encyclopaedia Britannica Online. Retrieved on 29 July 2009. Modern Standard Arabic is widely taught in schools, universities, and used in workplaces, government and the media.
Modern Standard Arabic derives from Classical Arabic, the only surviving member of the Old North Arabian dialect group, attested in Pre-Islamic Arabic inscriptions dating back to the 4th century. Versteegh, 1997, p. 33. Classical Arabic has also been a literary language and the liturgical language of Islam since its inception in the 7th century.
Arabic has lent many words to other languages of the Islamic world. During the Middle Ages, Arabic was a major vehicle of culture in Europe, especially in science, mathematics and philosophy. As a result, many European languages have also borrowed many words from it. Arabic influence is seen in Mediterranean languages, particularly Spanish, Portuguese, and Sicilian, due to both the proximity of European and Arab civilization and 700 years of Arab rule in the Iberian peninsula (see Al-Andalus).
Arabic has also borrowed words from many languages, including Hebrew, Persian and Syriac in early centuries, and contemporary European languages in modern times.
Arabic usually designates one of three main variants: Classical Arabic; Modern Standard Arabic; colloquial or dialectal Arabic.
Classical Arabic is the language found in the Qur'an and used from the period of Pre-Islamic Arabia to that of the Abbasid Caliphate. Classical Arabic is considered normative; modern authors attempt to follow the syntactic and grammatical norms laid down by classical grammarians (such as Sibawayh), and use the vocabulary defined in classical dictionaries (such as the LisÄn al-Arab).
Based on Classical Arabic, Modern Standard Arabic (ÙصØÙ fuṣḥÄ) is the literary language used in most current, printed Arabic publications, spoken by the Arabic media across North Africa and the Middle East, and understood by most educated Arabic speakers. "Literary Arabic" and "Standard Arabic" are less strictly defined terms that may refer to Modern Standard Arabic and/or Classical Arabic.
Colloquial or dialectal Arabic refers to the many national or regional varieties which constitute the everyday spoken language. Colloquial Arabic has many different regional variants; these sometimes differ enough to be mutually unintelligible and some linguists consider them distinct languages. "Arabic Language." Microsoft Encarta Online Encyclopedia 2009. Retrieved on 29 July 2009. The varieties are typically unwritten. They are often used in informal spoken media, such as soap operas and talk shows, as well as occasionally in certain forms of written media, such as poetry and printed advertising. The only variety of modern Arabic to have acquired official language status is Maltese, spoken in (predominately Roman Catholic) Malta and written with the Latin alphabet. It is descended from Classical Arabic through Siculo-Arabic and is not mutually intelligible with other varieties of Arabic. Most linguists list it as a separate language rather than as a dialect of Arabic.
The sociolinguistic situation of Arabic in modern times provides a prime example of the linguistic phenomenon of diglossia, which is the normal use of two separate varieties of the same language, usually in different social situations. In the case of Arabic, educated Arabs of any nationality can be assumed to speak both their local dialect and their school-taught Standard Arabic. When educated Arabs of different dialects engage in conversation (for example, a Moroccan speaking with a Lebanese), many speakers code-switch back and forth between the dialectal and standard varieties of the language, sometimes even within the same sentence. Arabic speakers often improve their familiarity with other dialects via music or film.
Like other languages, Modern Standard Arabic continues to evolve. Kaye, 1991. Many modern terms have entered into common usage, in some cases taken from other languages (for example, ÙÙÙÙ
film) or coined from existing lexical resources (for example, Ùات٠hÄtif "telephone" For these reasons, Modern Standard Arabic is generally treated separately in non-Arab sources.
The influence of Arabic has been most important in Islamic countries. Arabic is a major source of vocabulary for languages such as Amharic, Bengali, Berber, Catalan, Cypriot Greek, Gujarati, Hindustani , Indonesian, Kurdish, Malay, Marathi, Pashto, Persian, Portuguese, Punjabi, Rohingya, Sindhi, Spanish, Swahili, Tagalog, Turkish and Urdu as well as other languages in countries where these languages are spoken. For example, the Arabic word for book (/kitÄb/) has been borrowed in all the languages listed, with the exception of Spanish, Catalan and Portuguese which use the Latin-derived words "libro","llibre" and "livro", respectively, and Tagalog which uses "aklat". In addition, English has quite a few Arabic loan words, some directly but most through the medium of other Mediterranean languages. Other languages such as Maltese Maltese language - Britannica Online Encyclopedia and Kinubi derive from Arabic, rather than merely borrowing vocabulary or grammar rules.
The terms borrowed range from religious terminology (like Berber "prayer" Gregersen, 1977, p. 237.
Arabic was influenced by other languages as well. The most important sources of borrowings into (pre-Islamic) Arabic are Aramaic, which used to be the principal, international language of communication throughout the ancient Near and Middle East, Ethiopic, and to a lesser degree Hebrew (mainly religious concepts).
As Arabic occupied a position similar to Latin (in Europe) throughout the Islamic world many of the Arabic concepts in the field of science, philosophy, commerce etc., were often coined by non-native Arabic speakers, notably by Aramaic and Persian translators. This process of using Arabic roots in notably Turkish and Persian, to translate foreign concepts continued right until the 18th and 19th century, when large swaths of Arab-inhabited lands were under Ottoman rule.
Arabic is the language of the Qur'an. Arabic is often associated with Islam, but it is also spoken by Arab Christians, Mizrahi Jews and Iraqi Mandaeans.
Most of the world's Muslims do not speak Arabic as their native language but many can read the script and recite the words of religious texts. Some Muslim speakers of Arabic consider their language to be "the language chosen by God to speak to mankind" and is most notably understood by Muslims as being the lingua franca of the afterlife.
The earliest surviving texts in Proto-Arabic, or Ancient North Arabian, are the Hasaean inscriptions of eastern Saudi Arabia, from the 8th century BC, written not in the modern Arabic alphabet, nor in its Nabataean ancestor, but in variants of the epigraphic South Arabian musnad. These are followed by 6th-century BC Lihyanite texts from southeastern Saudi Arabia and the Thamudic texts found throughout Arabia and the Sinai, and not actually connected with Thamud. Later come the Safaitic inscriptions beginning in the 1st century BC, and the many Arabic personal names attested in Nabataean inscriptions (which are, however, written in Aramaic). From about the 2nd century BC, a few inscriptions from Qaryat al-Faw (near Sulayyil) reveal a dialect which is no longer considered "Proto-Arabic", but Pre-Classical Arabic.
By the fourth century AD, the Arab kingdoms of the Lakhmids in southern Iraq, the Ghassanids in southern Syria the Kindite Kingdom emerged in Central Arabia. Their courts were responsible for some notable examples of pre-Islamic Arabic poetry, and for some of the few surviving pre-Islamic Arabic inscriptions in the Arabic alphabet. A History of the Arabic Language
Colloquial Arabic is a collective term for the spoken varieties of Arabic used throughout the Arab world, which differ radically from the literary language. The main dialectal division is between the North African dialects and those of the Middle East, followed by that between sedentary dialects and the much more conservative Bedouin dialects. Speakers of some of these dialects are unable to converse with speakers of another dialect of Arabic. In particular, while Middle Easterners can generally understand one another, they often have trouble understanding North Africans (although the converse is not true, in part due to the popularity of Middle Easternâespecially Egyptianâfilms and other media).
One factor in the differentiation of the dialects is influence from the languages previously spoken in the areas, which have typically provided a significant number of new words, and have sometimes also influenced pronunciation or word order; however, a much more significant factor for most dialects is, as among Romance languages, retention (or change of meaning) of different classical forms. Thus Iraqi aku, Levantine fÄ«h, and North African kayÉn all mean "there is", and all come from Classical Arabic forms (yakÅ«n, fÄ«hi, kÄ'in respectively), but now sound very different.
Different Dialects of Arabic in the Arab World
The major dialect groups are:
* Egyptian Arabic, spoken by around 76 million in Egypt. It is one of the most understood varieties of Arabic. Closely related varieties are also spoken in Sudan.
* Gulf Arabic, spoken by around 34 million people in Arab states of the Persian Gulf and eastern Saudi Arabia.
* Iraqi Arabic, spoken by about 29 million people in Iraq. With significant differences between the Arabian-like dialects of the south and the more conservative dialects of the north. Closely related varieties are also spoken in Iran, Syria, and Turkey.
* North Mesopotamian Arabic, spoken by around 7 million people in northern Iraq, northern Syria and southern Turkey.
* Levantine Arabic, includes North Levantine Arabic, South Levantine Arabic, and Cypriot Arabic, and is spoken by almost 35 million people in Lebanon, Syria, Jordan, Palestine, Israel, Cyprus, and Turkey. It's also called Mediterranean Arabic.
* Maghrebi Arabic, heavily influenced by Berber in pronunciation, and includes Moroccan Arabic, Algerian Arabic, Algerian Saharan Arabic, Tunisian Arabic, and Libyan Arabic, and is spoken by around 45 million North Africans in Morocco, Western Sahara, Algeria, Tunisia, Libya, Niger, and western Egypt; it is mostly difficult for speakers of Near Eastern Arabic varieties to understand. The Berber influence in these dialects varies in degree. Kaplan and Baldauf, 2007, p. 48. See also Bateson, 2003, pp. 96-103 and Berber: Linguistic "Substratum" of North African Arabic by Ernest N. McCarus.
Other varieties include:
* Andalusi Arabic, spoken in Spain until 15th century, now extinct.
* Bahrani Arabic, spoken by Bahrani Shia in Bahrain, where it exhibits some differences from Bahraini Arabic. It is also spoken to a lesser extent in Oman.
* Central Asian Arabic, spoken in Uzbekistan, Tajikistan and Afghanistan, is highly endangered
* Hassaniya Arabic, spoken in Mauritania, some parts of Mali and Western Sahara
* Hejazi Arabic, spoken in Hejaz, western Saudi Arabia
* Judeo-Arabic dialects
* Maltese, spoken on the Mediterranean island of Malta, is the only one to have established itself as a fully separate language, with independent literary norms. In the course of its history the language has adopted numerous loanwords, phonetic and phonological features, and even some grammatical patterns, from Italian, Sicilian, and English. It is also the only Semitic tongue written in the Latin alphabet.
* Najdi Arabic, spoken in Nejd, central Saudi Arabia
* Shuwa Arabic, spoken in Chad, Cameroon, Niger, Nigeria, and Sudan
* Siculo Arabic, spoken on Sicily, South Italy until 14th century, developed into Maltese MED Magazine
* Sudanese Arabic, spoken in Sudan
* Yemeni Arabic, spoken in Yemen, southern Saudi Arabia, Djibouti, and Somalia
The phonemes below reflect the pronunciation of Modern Standard Arabic. There are minor variations from country to country. Additionally, these dialects can vary from region to region within a country.
Modern Standard Arabic has three vowels, with long and short forms of , , and . There are also two diphthongs: and .
See Arabic alphabet for explanations on the IPA phonetic symbols found in this chart.
# is pronounced by some speakers. This is especially characteristic of the Egyptian, Omani and some Yemeni dialects. In many parts of North Africa and in the Levant, it is pronounced .
# is pronounced only in , the name of God, q.e. Allah, when the word follows a, Ä, u or Å« (after i or Ä« it is unvelarized: bismi l-lÄh ).
# In many varieties, are actually epiglottal (despite what is reported in many earlier works).
# and are often post-velar though velar and uvular pronunciations are also possible.
Arabic has consonants traditionally termed "emphatic" exhibit simultaneous pharyngealization as well as varying degrees of velarization . This simultaneous articulation is described as "Retracted Tongue Root" by phonologists. e.g. In some transcription systems, emphasis is shown by capitalizing the letter, for example, is written â¹Dâº; in others the letter is underlined or has a dot below it, for example, .
Vowels and consonants can be phonologically short or long. Long (geminate) consonants are normally written doubled in Latin transcription (i.e. bb, dd, etc.), reflecting the presence of the Arabic diacritic mark shaddah, which indicates doubled consonants. In actual pronunciation, doubled consonants are held twice as long as short consonants. This consonant lengthening is phonemically contrastive: qabala "he accepted" vs. qabbala "he kissed."
Arabic has two kinds of syllables: open syllables (CV) and (CVV)âand closed syllables (CVC), (CVVC), and (CVCC), the latter two, which are (CVVC) and (CVCC) occuring only at the end of the sentence. Every syllable begins with a consonant. Syllables cannot begin with a vowel. Arabic phonology recognizes the glottal stop as an independent consonant, so in cases where a word begins with a vowel sound, as the definite article "al", for example, the word is recognized in Arabic as beginning with the consonant (glottal stop). When a word ends in a vowel and the following word begins with a glottal stop, then the glottal stop and the initial vowel of the word are in some cases elided, and the following consonant closes the final syllable of the preceding word, for example, baytu al-mudi:r "house (of) the director," which becomes .
Although word stress is not phonemically contrastive in Standard Arabic, it does bear a strong relationship to vowel length . The basic rules are:
* Only one of the last three syllables may be stressed.
* Given this restriction, the last "superheavy" syllable (containing a long vowel or ending in a consonant) is stressed.
* If there is no such syllable, the pre-final syllable is stressed if it is 'heavy.' Otherwise, the first allowable syllable is stressed.
* In Standard Arabic, a final long vowel may not be stressed. (This restriction does not apply to the spoken dialects, where original final long vowels have been shortened and secondary final long vowels have arisen.)
For example: ki-TAA-bun "book", KAA-ti-bun "writer", MAK-ta-bun "desk", ma-KAA-ti-bu "desks", mak-TA-ba-tun "library", KA-ta-buu (Modern Standard Arabic) "they wrote" = KA-ta-bu (dialect), ka-ta-BUU-hu (Modern Standard Arabic) "they wrote it" = ka-ta-BUU (dialect), ka-TA-ba-taa (Modern Standard Arabic) "they (dual, fem) wrote", ka-TAB-tu (Modern Standard Arabic) "I wrote" = ka-TABT (dialect). Doubled consonants count as two consonants: ma-JAL-la "magazine", ma-HALL "place".
Some dialects have different stress rules. In the Cairo (Egyptian Arabic) dialect, for example, a heavy syllable may not carry stress more than two syllables from the end of a word, hence mad-RA-sa "school", qaa-HI-ra "Cairo". In the Arabic of Sana, stress is often retracted: BAY-tayn "two houses", MAA-sat-hum "their table", ma-KAA-tiib "desks", ZAA-rat-hiin "sometimes", mad-RA-sat-hum "their school". (In this dialect, only syllables with long vowels or diphthongs are considered heavy; in a two-syllable word, the final syllable can be stressed only if the preceding syllable is light; and in longer words, the final syllable cannot be stressed.)
In some dialects, there may be more or fewer phonemes than those listed in the chart above. For example, non-Arabic is used in the Maghrebi dialects as well in the written language mostly for foreign names. Semitic became extremely early on in Arabic before it was written down; a few modern Arabic dialects, such as Iraqi (influenced by Persian and Turkish) distinguish between and .
Interdental fricatives ( and ) are rendered as stops and in some dialects (such as Egyptian, Levantine, and much of the Maghreb); some of these dialects render them as and in "learned" words from the Standard language. Early in the expansion of Arabic, the separate emphatic phonemes and coallesced into a single phoneme, becoming one or the other. Predictably, dialects without interdental fricatives use exclusively, while dialects with such fricatives use . Again, in "learned" words from the Standard language, is rendered as (in Egypt & the Levant) or (in North Africa) in dialects without interdental fricatives.
Another key distinguishing mark of Arabic dialects is how they render the original velar and uvular stops , (Proto-Semitic ), and :
* Ù retains its original pronunciation in widely scattered regions such as Yemen, Morocco, and urban areas of the Maghreb. It is pronounced as a glottal stop in several prestige dialects, such as those spoken in Cairo, Beirut and Damascus. But it is rendered as a voiced velar stop in Gulf Arabic, Iraqi Arabic, Upper Egypt, much of the Maghreb, and less urban parts of the Levant (e.g. Jordan). Some traditionally Christian villages in rural areas of the Levant render the sound as , as do Shia Bahrainis. In some Gulf dialects, it is palatalized to or . It is pronounced as a voiced uvular constrictive in Sudanese Arabic. Many dialects with a modified pronunciation for maintain the pronunciation in certain words (often with religious or educational overtones) borrowed from the Classical language.
* ج retains its pronunciation in Iraq and much of the Arabian Peninsula, but is pronounced in most of North Egypt and parts of Yemen, in Morocco and the Levant, and in some words in much of Gulf Arabic.
* Ù usually retains its original pronunciation, but is palatalized to in many words in Palestine, Iraq and much of the Arabian Peninsula. Often a distinction is made between the suffixes (you, masc.) and (you, fem.), which become and , respectively. In Sana Arabic, is pronounced .
Visualization of Arabic grammar from the Quranic Arabic Corpus
Compared with other Semitic language systems, Classical Arabic is distinguished by, "its almost (too perfect) algebraic-looking grammar, i.e. root pattern and morphology." Hetzron, 1997, p. 229. Nouns in Literary Arabic have three grammatical cases (nominative, accusative, and genitive [also used when the noun is governed by a preposition]); three numbers (singular, dual and plural); two genders (masculine and feminine); and three "states" (indefinite, definite, and construct). The cases of singular nouns (other than those that end in long Ä) are indicated by suffixed short vowels (/-u/ for nominative, /-a/ for accusative, /-i/ for genitive). The feminine singular is often marked by /-at/, which is reduced to /-ah/ or /-a/ before a pause. Plural is indicated either through endings (the sound plural) or internal modification (the broken plural). Definite nouns include all proper nouns, all nouns in "construct state" and all nouns which are prefixed by the definite article /al-/. Indefinite singular nouns (other than those that end in long Ä) add a final /-n/ to the case-marking vowels, giving /-un/, /-an/ or /-in/ (which is also referred to as nunation or tanwÄ«n).
Verbs in Literary Arabic are marked for person (first, second, or third), gender, and number. They are conjugated in two major paradigms (termed perfective and imperfective, or past and non-past); two voices (active and passive); and five moods in the imperfective (indicative, imperative, subjunctive, jussive and energetic). There are also two participles (active and passive) and a verbal noun, but no infinitive. As indicated by the differing terms for the two tense systems, there is some disagreement over whether the distinction between the two systems should be most accurately characterized as tense, aspect or a combination of the two. The perfective aspect is constructed using fused suffixes that combine person, number and gender in a single morpheme, while the imperfective aspect is constructed using a combination of prefixes (primarily encoding person) and suffixes (primarily encoding gender and number). The moods other than imperative are primarily marked by suffixes (/u/ for indicative, /a/ for subjunctive, no ending for jussive, /an/ for energetic). The imperative has the endings of the jussive but lacks any prefixes. The passive is marked through internal vowel changes. Plural forms for the verb are only used when the subject is not mentioned, or is preceding it, and the feminine singular is used for all non-human plurals.
Adjectives in Literary Arabic are marked for case, number, gender and state, as for nouns. However, the plural of all non-human nouns is always combined with a singular feminine adjective, which takes the /-ah/ or /-at/ suffix.
Pronouns in Literary Arabic are marked for person, number and gender. There are two varieties, independent pronouns and enclitics. Enclitic pronouns are attached to the end of a verb, noun or preposition and indicate verbal and prepositional objects or possession of nouns. The first-person singular pronoun has a different enclitic form used for verbs (/-ni/) and for nouns or prepositions (/-Ä«/ after consonants, /-ya/ after vowels).
Nouns, verbs, pronouns and adjectives agree with each other in all respects. However, non-human plural nouns are grammatically considered to be feminine singular. Furthermore, a verb in a verb-initial sentence is marked as singular regardless of its semantic number when the subject of the verb is explicitly mentioned as a noun. Numerals between three and ten show "chiasmic" agreement, in that grammatically masculine numerals have feminine marking and vice versa.
The spoken dialects have lost the case distinctions and make only limited use of the dual (it occurs only on nouns and its use is no longer required in all circumstances). They have lost the mood distinctions other than imperative, but many have since gained new moods through the use of prefixes (most often /bi-/ for indicative vs. unmarked subjunctive). They have also mostly lost the indefinite "nunation" and the internal passive. Modern Standard Arabic maintains the grammatical distinctions of Literary Arabic except that the energetic mood is almost never used; in addition, Modern Standard Arabic sometimes drop the final short vowels that indicate case and mood.
As in many other Semitic languages, Arabic verb formation is based on a (usually) triconsonantal root, which is not a word in itself but contains the semantic core. The consonants , for example, indicate write, indicate read, indicate eat, etc. Words are formed by supplying the root with a vowel structure and with affixes. (Traditionally, Arabic grammarians have used the root , do, as a template to discuss word formation.) From any particular root, up to fifteen different verbs can be formed, each with its own template; these are referred to by Western scholars as "form I", "form II", and so on through "form XV". These forms, and their associated participles and verbal nouns, are the primary means of forming vocabulary in Arabic. Forms XI to XV are incidental.
An example of a text written in Arabic calligraphy.
The Arabic alphabet derives from the Aramaic script through Nabatean, to which it bears a loose resemblance like that of Coptic or Cyrillic script to Greek script. Traditionally, there were several differences between the Western (North African) and Middle Eastern version of the alphabetâin particular, the fa and qaf had a dot underneath and a single dot above respectively in the Maghreb, and the order of the letters was slightly different (at least when they were used as numerals). However, the old Maghrebi variant has been abandoned except for calligraphic purposes in the Maghreb itself, and remains in use mainly in the Quranic schools (zaouias) of West Africa. Arabic, like all other Semitic languages (except for the Latin-written Maltese, and the languages with the Ge'ez script), is written from right to left. There are several styles of script, notably Naskh which is used in print and by computers, and Ruq'ah which is commonly used in handwriting. Hanna, 1972, p. 2
After the definitive fixing of the Arabic script around 786, by Khalil ibn Ahmad al Farahidi, many styles were developed, both for the writing down of the Qur'an and other books, and for inscriptions on monuments as decoration.
Arabic calligraphy has not fallen out of use as calligraphy has in the Western world, and is still considered by Arabs as a major art form; calligraphers are held in great esteem. Being cursive by nature, unlike the Latin alphabet, Arabic script is used to write down a verse of the Qur'an, a Hadith, or simply a proverb, in a spectacular composition. The composition is often abstract, but sometimes the writing is shaped into an actual form such as that of an animal. One of the current masters of the genre is Hassan Massoudy
There are a number of different standards of Arabic transliteration: methods of accurately and efficiently representing Arabic with the Latin alphabet. There are multiple conflicting motivations for transliteration. Scholarly systems are intended to accurately and unambiguously represent the phonemes of Arabic, generally making the phonetics more explicit than the original word in the Arabic alphabet. These systems are heavily reliant on diacritical marks such as "š" for the sound equivalently written sh in English. In some cases, the sh or kh sounds can be represented by italicizing or underlining them that way, they can be distinguished from separate s and h sounds or k and h sounds, respectively. (Compare gashouse to gash.) At first sight, this may be difficult to recognize. Less scientific systems often use digraphs (like sh and kh), which are usually more simple to read, but sacrifice the definiteness of the scientific systems. Such systems may be intended to help readers who are neither Arabic speakers nor linguists to intuitively pronounce Arabic names and phrases. An example of such a system is the Bahá'à orthography. A third type of transliteration seeks to represent an equivalent of the Arabic spelling with Latin letters, for use by Arabic speakers when Arabic writing is not available (for example, when using an ASCII communication device).
An example is the system used by the US military, Standard Arabic Technical Transliteration System or SATTS, which represents each Arabic letter with a unique symbol in the ASCII range to provide a one-to-one mapping from Arabic to ASCII and back. This system, while facilitating typing on English keyboards, presents its own ambiguities and disadvantages. During the last few decades and especially since the 1990s, Western-invented text communication technologies have become prevalent in the Arab world, such as personal computers, the World Wide Web, email, Bulletin board systems, IRC, instant messaging and mobile phone text messaging. Most of these technologies originally had the ability to communicate using the Latin alphabet only, and some of them still do not have the Arabic alphabet as an optional feature. As a result, Arabic speaking users communicated in these technologies by transliterating the Arabic text using the Latin script, sometimes known as IM Arabic.
To handle those Arabic letters that cannot be accurately represented using the Latin script, numerals and other characters were appropriated. For example, the numeral "3" may be used to represent the Arabic letter "ع", ayn. There is no universal name for this type of transliteration, but some have named it Arabic Chat Alphabet. Other systems of transliteration exist, such as using dots or capitalization to represent the "emphatic" counterparts of certain consonants. For instance, using capitalization, the letter "د", or daal, may be represented by d. Its emphatic counterpart, "ض", may be written as D.
In most of present-day North Africa, the Western Arabic numerals (0, 1, 2, 3, 4, 5, 6, 7, 8, 9) are used. However in Egypt and Arabic-speaking countries to the east of it, the Eastern Arabic numerals ( ) are in use. When representing a number in Arabic, the lowest-valued position is placed on the right, so the order of positions is the same as in left-to-right scripts. Sequences of digits such as telephone numbers are read from left to right, but numbers are spoken in the traditional Arabic fashion, with units and tens reversed from the modern English usage. For example, 24 is said "four and twenty", and 1975 is said "one thousand and nine hundred and five and seventy."
Academy of the Arabic Language is the name of a number of language-regulation bodies formed in Arab countries. The most active are in Damascus and Cairo. They review language development, monitor new words and approve inclusion of new words into their published standard dictionaries. They also publish old and historical Arabic manuscripts.
Because the Quran is written in Arabic and all Islamic terms are in Arabic, millions of Muslims (both Arab and non-Arab) study the language. Arabic has been taught in many elementary and secondary schools, especially Muslim schools, worldwide. Universities around the world have classes teaching Arabic as part of their foreign languages, Middle Eastern studies, religious studies courses. Arabic language schools exist to assist students in learning Arabic outside of the academic world. Many Arabic language schools are located in the Arab world and other Muslim countries. Software and books with tapes are also important part of Arabic learning, as many of Arabic learners may live in places where there are no academic or Arabic language school classes available. Radio series of Arabic language classes are also provided from some radio stations. A number of websites on the Internet provide online classes for all levels as a means of distance education.
* Arabic alphabet
* Arabic calligraphy
* Arabic diglossia
* Arabic influence on Spanish
* Arabic literature
* Arabist
* Dictionary of Modern Written Arabic
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* List of Arabic loanwords in English
* List of French words of Arabic origin
* List of Islamic terms in Arabic
* List of Portuguese words of Arabic origin
* List of replaced loanwords in Turkish
* Literary Arabic
* Macrolanguage
* Varieties of Arabic
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* Journal of Arabic and Islamic Studies
* Google Ta3reeb - Arabic Keyboard using English/Latin Characters
* eiktub - realtime Arabic transliteration
* Lane's Arabic-English Lexicon, an 8-volume, 3000-page dictionary available for download in PDF format.
* Learn Classical Arabic Online
* Arabic - a Category III language Languages which are exceptionally difficult for native English speakers
* Yalla-2009, a free software to understand how Arabic language is built
* Arabic grammar online
* Arabic language pronunciation applet with audio samples
* The Expansion of the Arabic language video on YouTube
* Free Arabic Course Online
* Dr. Habash's Introduction to Arabic Natural Language Processing
* The Arabic Language & the Qur'an
* Software utility for converting between Arabic language
* Software for Arabic language
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