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Various species intercalate into hexagonal BN, such as intercalate or alkali metals.
Preparation of cubic BN
c-BN is prepared analogously to the preparation of synthetic diamond from graphite. Direct conversion of hexagonal boron nitride to the cubic form has been observed at pressures between 5 and 18 GPa and temperatures between 1730 and 3230 °C, that is similar parameters as for direct graphite-diamond conversion. The addition of a small amount of boron oxide can lower the required pressure to 4–7 GPa and temperature to 1500 °C. As in diamond synthesis, to further reduce the conversion pressures and temperatures, a catalyst is added, such as lithium, potassium, or magnesium, their nitrides, their fluoronitrides, water with ammonium compounds, or hydrazine. Other industrial synthesis methods, again borrowed from diamond growth, use crystal growth in a temperature gradient, or explosive shock wave. The shock wave method is used to produce material called heterodiamond, a superhard compound of boron, carbon, and nitrogen.
Low-pressure deposition of thin films of cubic boron nitride is possible. As in diamond growth, the major problem is to suppress the growth of hexagonal phases (h-BN or graphite, respectively). Whereas in diamond growth this is achieved by adding hydrogen gas, boron trifluoride is used for c-BN. Ion beam deposition, plasma-enhanced chemical vapor deposition, pulsed laser deposition, reactive sputtering, and other physical vapor deposition methods are used as well.
Preparation of wurtzite BN
Wurtzite BN can be obtained via static high-pressure or dynamic shock methods. The limits of its stability are not well defined. Both c-BN and w-BN are formed by compressing h-BN, but formation of w-BN occurs at much lower temperatures close to 1700 °C. | Boron nitride | Wikipedia | 389 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Production statistics
Whereas the production and consumption figures for the raw materials used for BN synthesis, namely boric acid and boron trioxide, are well known (see boron), the corresponding numbers for the boron nitride are not listed in statistical reports. An estimate for the 1999 world production is 300 to 350 metric tons. The major producers and consumers of BN are located in the United States, Japan, China and Germany. In 2000, prices varied from about $75–120/kg for standard industrial-quality h-BN and were about up to $200–400/kg for high purity BN grades.
Applications
Hexagonal BN
Hexagonal BN (h-BN) is the most widely used polymorph. It is a good lubricant at both low and high temperatures (up to 900 °C, even in an oxidizing atmosphere). h-BN lubricant is particularly useful when the electrical conductivity or chemical reactivity of graphite (alternative lubricant) would be problematic. In internal combustion engines, where graphite could be oxidized and turn into carbon sludge, h-BN with its superior thermal stability can be added to engine lubricants. As with all nano-particle suspensions, Brownian-motion settlement is a problem. Settlement can clog engine oil filters, which limits solid lubricant applications in a combustion engine to automotive racing, where engine re-building is common. Since carbon has appreciable solubility in certain alloys (such as steels), which may lead to degradation of properties, BN is often superior for high temperature and/or high pressure applications. Another advantage of h-BN over graphite is that its lubricity does not require water or gas molecules trapped between the layers. Therefore, h-BN lubricants can be used in vacuum, such as space applications. The lubricating properties of fine-grained h-BN are used in cosmetics, paints, dental cements, and pencil leads.
Hexagonal BN was first used in cosmetics around 1940 in Japan. Because of its high price, h-BN was abandoned for this application. Its use was revitalized in the late 1990s with the optimization h-BN production processes, and currently h-BN is used by nearly all leading producers of cosmetic products for foundations, make-up, eye shadows, blushers, kohl pencils, lipsticks and other skincare products. | Boron nitride | Wikipedia | 505 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Because of its excellent thermal and chemical stability, boron nitride ceramics and coatings are used high-temperature equipment. h-BN can be included in ceramics, alloys, resins, plastics, rubbers, and other materials, giving them self-lubricating properties. Such materials are suitable for construction of e.g. bearings and in steelmaking. Many quantum devices use multilayer h-BN as a substrate material. It can also be used as a dielectric in resistive random access memories.
Hexagonal BN is used in xerographic process and laser printers as a charge leakage barrier layer of the photo drum. In the automotive industry, h-BN mixed with a binder (boron oxide) is used for sealing oxygen sensors, which provide feedback for adjusting fuel flow. The binder utilizes the unique temperature stability and insulating properties of h-BN.
Parts can be made by hot pressing from four commercial grades of h-BN. Grade HBN contains a boron oxide binder; it is usable up to 550–850 °C in oxidizing atmosphere and up to 1600 °C in vacuum, but due to the boron oxide content is sensitive to water. Grade HBR uses a calcium borate binder and is usable at 1600 °C. Grades HBC and HBT contain no binder and can be used up to 3000 °C.
Boron nitride nanosheets (h-BN) can be deposited by catalytic decomposition of borazine at a temperature ~1100 °C in a chemical vapor deposition setup, over areas up to about 10 cm2. Owing to their hexagonal atomic structure, small lattice mismatch with graphene (~2%), and high uniformity they are used as substrates for graphene-based devices. BN nanosheets are also excellent proton conductors. Their high proton transport rate, combined with the high electrical resistance, may lead to applications in fuel cells and water electrolysis.
h-BN has been used since the mid-2000s as a bullet and bore lubricant in precision target rifle applications as an alternative to molybdenum disulfide coating, commonly referred to as "moly". It is claimed to increase effective barrel life, increase intervals between bore cleaning and decrease the deviation in point of impact between clean bore first shots and subsequent shots. | Boron nitride | Wikipedia | 479 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
h-BN is used as a release agent in molten metal and glass applications. For example, ZYP Coatings developed and currently produces a line of paintable h-BN coatings that are used by manufacturers of molten aluminium, non-ferrous metal, and glass. Because h-BN is nonwetting and lubricious to these molten materials, the coated surface (i.e. mold or crucible) does not stick to the material.
Cubic BN
Cubic boron nitride (CBN or c-BN) is widely used as an abrasive. Its usefulness arises from its insolubility in iron, nickel, and related alloys at high temperatures, whereas diamond is soluble in these metals. Polycrystalline c-BN (PCBN) abrasives are therefore used for machining steel, whereas diamond abrasives are preferred for aluminum alloys, ceramics, and stone. When in contact with oxygen at high temperatures, BN forms a passivation layer of boron oxide. Boron nitride binds well with metals due to formation of interlayers of metal borides or nitrides. Materials with cubic boron nitride crystals are often used in the tool bits of cutting tools. For grinding applications, softer binders such as resin, porous ceramics and soft metals are used. Ceramic binders can be used as well. Commercial products are known under names "Borazon" (by Hyperion Materials & Technologies), and "Elbor" or "Cubonite" (by Russian vendors).
Contrary to diamond, large c-BN pellets can be produced in a simple process (called sintering) of annealing c-BN powders in nitrogen flow at temperatures slightly below the BN decomposition temperature. This ability of c-BN and h-BN powders to fuse allows cheap production of large BN parts.
Similar to diamond, the combination in c-BN of highest thermal conductivity and electrical resistivity is ideal for heat spreaders.
As cubic boron nitride consists of light atoms and is very robust chemically and mechanically, it is one of the popular materials for X-ray membranes: low mass results in small X-ray absorption, and good mechanical properties allow usage of thin membranes, further reducing the absorption. | Boron nitride | Wikipedia | 467 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Amorphous BN
Layers of amorphous boron nitride (a-BN) are used in some semiconductor devices, e.g. MOSFETs. They can be prepared by chemical decomposition of trichloroborazine with caesium, or by thermal chemical vapor deposition methods. Thermal CVD can be also used for deposition of h-BN layers, or at high temperatures, c-BN.
Other forms of boron nitride
Atomically thin boron nitride
Hexagonal boron nitride can be exfoliated to mono or few atomic layer sheets. Due to its analogous structure to that of graphene, atomically thin boron nitride is sometimes called white graphene.
Mechanical properties
Atomically thin boron nitride is one of the strongest electrically insulating materials. Monolayer boron nitride has an average Young's modulus of 0.865TPa and fracture strength of 70.5GPa, and in contrast to graphene, whose strength decreases dramatically with increased thickness, few-layer boron nitride sheets have a strength similar to that of monolayer boron nitride.
Thermal conductivity
Atomically thin boron nitride has one of the highest thermal conductivity coefficients (751 W/mK at room temperature) among semiconductors and electrical insulators, and its thermal conductivity increases with reduced thickness due to less intra-layer coupling.
Thermal stability
The air stability of graphene shows a clear thickness dependence: monolayer graphene is reactive to oxygen at 250 °C, strongly doped at 300 °C, and etched at 450 °C; in contrast, bulk graphite is not oxidized until 800 °C. Atomically thin boron nitride has much better oxidation resistance than graphene. Monolayer boron nitride is not oxidized till 700 °C and can sustain up to 850 °C in air; bilayer and trilayer boron nitride nanosheets have slightly higher oxidation starting temperatures. The excellent thermal stability, high impermeability to gas and liquid, and electrical insulation make atomically thin boron nitride potential coating materials for preventing surface oxidation and corrosion of metals and other two-dimensional (2D) materials, such as black phosphorus. | Boron nitride | Wikipedia | 465 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Better surface adsorption
Atomically thin boron nitride has been found to have better surface adsorption capabilities than bulk hexagonal boron nitride. According to theoretical and experimental studies, atomically thin boron nitride as an adsorbent experiences conformational changes upon surface adsorption of molecules, increasing adsorption energy and efficiency. The synergic effect of the atomic thickness, high flexibility, stronger surface adsorption capability, electrical insulation, impermeability, high thermal and chemical stability of BN nanosheets can increase the Raman sensitivity by up to two orders, and in the meantime attain long-term stability and reusability not readily achievable by other materials.
Dielectric properties
Atomically thin hexagonal boron nitride is an excellent dielectric substrate for graphene, molybdenum disulfide (), and many other 2D material-based electronic and photonic devices. As shown by electric force microscopy (EFM) studies, the electric field screening in atomically thin boron nitride shows a weak dependence on thickness, which is in line with the smooth decay of electric field inside few-layer boron nitride revealed by the first-principles calculations.
Raman characteristics
Raman spectroscopy has been a useful tool to study a variety of 2D materials, and the Raman signature of high-quality atomically thin boron nitride was first reported by Gorbachev et al. in 2011. and Li et al. However, the two reported Raman results of monolayer boron nitride did not agree with each other. Cai et al., therefore, conducted systematic experimental and theoretical studies to reveal the intrinsic Raman spectrum of atomically thin boron nitride. It reveals that atomically thin boron nitride without interaction with a substrate has a G band frequency similar to that of bulk hexagonal boron nitride, but strain induced by the substrate can cause Raman shifts. Nevertheless, the Raman intensity of G band of atomically thin boron nitride can be used to estimate layer thickness and sample quality.
Boron nitride nanomesh | Boron nitride | Wikipedia | 452 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Boron nitride nanomesh is a nanostructured two-dimensional material. It consists of a single BN layer, which forms by self-assembly a highly regular mesh after high-temperature exposure of a clean rhodium or ruthenium surface to borazine under ultra-high vacuum. The nanomesh looks like an assembly of hexagonal pores. The distance between two pore centers is 3.2 nm and the pore diameter is ~2 nm. Other terms for this material are boronitrene or white graphene.
The boron nitride nanomesh is air-stable and compatible with some liquids. up to temperatures of 800 °C.
Boron nitride nanotubes
Boron nitride tubules were first made in 1989 by Shore and Dolan This work was patented in 1989 and published in 1989 thesis (Dolan) and then 1993 Science. The 1989 work was also the first preparation of amorphous BN by B-trichloroborazine and cesium metal.
Boron nitride nanotubes were predicted in 1994 and experimentally discovered in 1995. They can be imagined as a rolled up sheet of h-boron nitride. Structurally, it is a close analog of the carbon nanotube, namely a long cylinder with diameter of several to hundred nanometers and length of many micrometers, except carbon atoms are alternately substituted by nitrogen and boron atoms. However, the properties of BN nanotubes are very different: whereas carbon nanotubes can be metallic or semiconducting depending on the rolling direction and radius, a BN nanotube is an electrical insulator with a bandgap of ~5.5 eV, basically independent of tube chirality and morphology. In addition, a layered BN structure is much more thermally and chemically stable than a graphitic carbon structure.
Boron nitride aerogel | Boron nitride | Wikipedia | 395 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Boron nitride aerogel is an aerogel made of highly porous BN. It typically consists of a mixture of deformed BN nanotubes and nanosheets. It can have a density as low as 0.6 mg/cm3 and a specific surface area as high as 1050 m2/g, and therefore has potential applications as an absorbent, catalyst support and gas storage medium. BN aerogels are highly hydrophobic and can absorb up to 160 times their weight in oil. They are resistant to oxidation in air at temperatures up to 1200 °C, and hence can be reused after the absorbed oil is burned out by flame. BN aerogels can be prepared by template-assisted chemical vapor deposition using borazine as the feed gas.
Composites containing BN
Addition of boron nitride to silicon nitride ceramics improves the thermal shock resistance of the resulting material. For the same purpose, BN is added also to silicon nitride-alumina and titanium nitride-alumina ceramics. Other materials being reinforced with BN include alumina and zirconia, borosilicate glasses, glass ceramics, enamels, and composite ceramics with titanium boride-boron nitride, titanium boride-aluminium nitride-boron nitride, and silicon carbide-boron nitride composition.
Zirconia Stabilized Boron Nitride (ZSBN) is produced by adding zirconia to BN, enhancing its thermal shock resistance and mechanical strength through a sintering process. It offers better performance characteristics including Superior corrosion and erosion resistance over a wide temperature range. Its unique combination of thermal conductivity, lubricity, mechanical strength, and stability makes it suitable for various applications including cutting tools and wear-resistant coatings, thermal and electrical insulation, aerospace and defense, and high-temperature components.
Pyrolytic boron nitride (PBN)
Pyrolytic boron nitride (PBN), also known as Chemical vapour-deposited Boron Nitride(CVD-BN), is a high-purity ceramic material characterized by exceptional chemical resistance and mechanical strength at high temperatures.
Pyrolytic boron nitride is typically prepared through the thermal decomposition of boron trichloride and ammonia vapors on graphite substrates at 1900°C. | Boron nitride | Wikipedia | 489 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Pyrolytic boron nitride (PBN) generally has a hexagonal structure similar to hexagonal boron nitride (hBN), though it can exhibit stacking faults or deviations from the ideal lattice. Pyrolytic boron nitride (PBN) shows some remarkable attributes, including exceptional chemical inertness, high dielectric strength, excellent thermal shock resistance, non-wettability, non-toxicity, oxidation resistance, and minimal outgassing.
Due to a highly ordered planar texture similar to pyrolytic graphite (PG), it exhibits anisotropic properties such as lower dielectric constant vertical to the crystal plane and higher bending strength along the crystal plane. PBN material has been widely manufactured as crucibles of compound semiconductor crystals, output windows and dielectric rods of traveling-wave tubes, high-temperature jigs and insulator.
Health issues
Boron nitride (along with , NbN, and BNC) is generally considered to be non-toxic and does not exhibit chemical activity in biological systems. Due to its excellent safety profile and lubricious properties, boron nitride finds widespread use in various applications, including cosmetics and food processing equipment. | Boron nitride | Wikipedia | 258 | 3370 | https://en.wikipedia.org/wiki/Boron%20nitride | Physical sciences | Ceramic compounds | Chemistry |
Beryllium is a chemical element; it has symbol Be and atomic number 4. It is a steel-gray, hard, strong, lightweight and brittle alkaline earth metal. It is a divalent element that occurs naturally only in combination with other elements to form minerals. Gemstones high in beryllium include beryl (aquamarine, emerald, red beryl) and chrysoberyl. It is a relatively rare element in the universe, usually occurring as a product of the spallation of larger atomic nuclei that have collided with cosmic rays. Within the cores of stars, beryllium is depleted as it is fused into heavier elements. Beryllium constitutes about 0.0004 percent by mass of Earth's crust. The world's annual beryllium production of 220 tons is usually manufactured by extraction from the mineral beryl, a difficult process because beryllium bonds strongly to oxygen.
In structural applications, the combination of high flexural rigidity, thermal stability, thermal conductivity and low density (1.85 times that of water) make beryllium a desirable aerospace material for aircraft components, missiles, spacecraft, and satellites. Because of its low density and atomic mass, beryllium is relatively transparent to X-rays and other forms of ionizing radiation; therefore, it is the most common window material for X-ray equipment and components of particle detectors. When added as an alloying element to aluminium, copper (notably the alloy beryllium copper), iron, or nickel, beryllium improves many physical properties. For example, tools and components made of beryllium copper alloys are strong and hard and do not create sparks when they strike a steel surface. In air, the surface of beryllium oxidizes readily at room temperature to form a passivation layer 1–10 nm thick that protects it from further oxidation and corrosion. The metal oxidizes in bulk (beyond the passivation layer) when heated above , and burns brilliantly when heated to about .
The commercial use of beryllium requires the use of appropriate dust control equipment and industrial controls at all times because of the toxicity of inhaled beryllium-containing dusts that can cause a chronic life-threatening allergic disease, berylliosis, in some people. Berylliosis is typically manifested by chronic pulmonary fibrosis and, in severe cases, right sided heart failure and death.
Characteristics | Beryllium | Wikipedia | 488 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Physical properties
Beryllium is a steel gray and hard metal that is brittle at room temperature and has a close-packed hexagonal crystal structure. It has exceptional stiffness (Young's modulus 287 GPa) and a melting point of 1287 °C. The modulus of elasticity of beryllium is approximately 35% greater than that of steel. The combination of this modulus and a relatively low density results in an unusually fast sound conduction speed in beryllium – about 12.9 km/s at ambient conditions. Other significant properties are high specific heat () and thermal conductivity (), which make beryllium the metal with the best heat dissipation characteristics per unit weight. In combination with the relatively low coefficient of linear thermal expansion (11.4 × 10−6 K−1), these characteristics result in a unique stability under conditions of thermal loading.
Nuclear properties
Naturally occurring beryllium, save for slight contamination by the cosmogenic radioisotopes, is isotopically pure beryllium-9, which has a nuclear spin of . Beryllium has a large scattering cross section for high-energy neutrons, about 6 barns for energies above approximately 10 keV. Therefore, it works as a neutron reflector and neutron moderator, effectively slowing the neutrons to the thermal energy range of below 0.03 eV, where the total cross section is at least an order of magnitude lower; the exact value strongly depends on the purity and size of the crystallites in the material.
The single primordial beryllium isotope 9Be also undergoes a (n,2n) neutron reaction with neutron energies over about 1.9 MeV, to produce 8Be, which almost immediately breaks into two alpha particles. Thus, for high-energy neutrons, beryllium is a neutron multiplier, releasing more neutrons than it absorbs. This nuclear reaction is:
+ n → 2 + 2 n
Neutrons are liberated when beryllium nuclei are struck by energetic alpha particles producing the nuclear reaction
+ → + n
where is an alpha particle and is a carbon-12 nucleus.
Beryllium also releases neutrons under bombardment by gamma rays. Thus, natural beryllium bombarded either by alphas or gammas from a suitable radioisotope is a key component of most radioisotope-powered nuclear reaction neutron sources for the laboratory production of free neutrons. | Beryllium | Wikipedia | 495 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Small amounts of tritium are liberated when nuclei absorb low energy neutrons in the three-step nuclear reaction
+ n → + , → + β−, + n → +
has a half-life of only 0.8 seconds, β− is an electron, and has a high neutron absorption cross section. Tritium is a radioisotope of concern in nuclear reactor waste streams.
Optical properties
As a metal, beryllium is transparent or translucent to most wavelengths of X-rays and gamma rays, making it useful for the output windows of X-ray tubes and other such apparatus.
Isotopes and nucleosynthesis
Both stable and unstable isotopes of beryllium are created in stars, but the radioisotopes do not last long. It is believed that most of the stable beryllium in the universe was originally created in the interstellar medium when cosmic rays induced fission in heavier elements found in interstellar gas and dust. Primordial beryllium contains only one stable isotope, 9Be, and therefore beryllium is, uniquely among all stable elements with an even atomic number, a monoisotopic and mononuclidic element.
Radioactive cosmogenic 10Be is produced in the atmosphere of the Earth by the cosmic ray spallation of oxygen. 10Be accumulates at the soil surface, where its relatively long half-life (1.36 million years) permits a long residence time before decaying to boron-10. Thus, 10Be and its daughter products are used to examine natural soil erosion, soil formation and the development of lateritic soils, and as a proxy for measurement of the variations in solar activity and the age of ice cores. The production of 10Be is inversely proportional to solar activity, because increased solar wind during periods of high solar activity decreases the flux of galactic cosmic rays that reach the Earth. Nuclear explosions also form 10Be by the reaction of fast neutrons with 13C in the carbon dioxide in air. This is one of the indicators of past activity at nuclear weapon test sites.
The isotope 7Be (half-life 53 days) is also cosmogenic, and shows an atmospheric abundance linked to sunspots, much like 10Be. | Beryllium | Wikipedia | 448 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
8Be has a very short half-life of about 8 s that contributes to its significant cosmological role, as elements heavier than beryllium could not have been produced by nuclear fusion in the Big Bang. This is due to the lack of sufficient time during the Big Bang's nucleosynthesis phase to produce carbon by the fusion of 4He nuclei and the very low concentrations of available beryllium-8. British astronomer Sir Fred Hoyle first showed that the energy levels of 8Be and 12C allow carbon production by the so-called triple-alpha process in helium-fueled stars where more nucleosynthesis time is available. This process allows carbon to be produced in stars, but not in the Big Bang. Star-created carbon (the basis of carbon-based life) is thus a component in the elements in the gas and dust ejected by AGB stars and supernovae (see also Big Bang nucleosynthesis), as well as the creation of all other elements with atomic numbers larger than that of carbon.
The 2s electrons of beryllium may contribute to chemical bonding. Therefore, when 7Be decays by L-electron capture, it does so by taking electrons from its atomic orbitals that may be participating in bonding. This makes its decay rate dependent to a measurable degree upon its chemical surroundings – a rare occurrence in nuclear decay.
The shortest-lived known isotope of beryllium is 16Be, which decays through neutron emission with a half-life of . The exotic isotopes 11Be and 14Be are known to exhibit a nuclear halo. This phenomenon can be understood as the nuclei of 11Be and 14Be have, respectively, 1 and 4 neutrons orbiting substantially outside the classical Fermi 'waterdrop' model of the nucleus.
Occurrence
The Sun has a concentration of 0.1 parts per billion (ppb) of beryllium. Beryllium has a concentration of 2 to 6 parts per million (ppm) in the Earth's crust and is the 47th most abundant element. It is most concentrated in the soils at 6 ppm. Trace amounts of 9Be are found in the Earth's atmosphere. The concentration of beryllium in sea water is 0.2–0.6 parts per trillion. In stream water, however, beryllium is more abundant with a concentration of 0.1 ppb. | Beryllium | Wikipedia | 487 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Beryllium is found in over 100 minerals, but most are uncommon to rare. The more common beryllium containing minerals include: bertrandite (Be4Si2O7(OH)2), beryl (Al2Be3Si6O18), chrysoberyl (Al2BeO4) and phenakite (Be2SiO4). Precious forms of beryl are aquamarine, red beryl and emerald.
The green color in gem-quality forms of beryl comes from varying amounts of chromium (about 2% for emerald).
The two main ores of beryllium, beryl and bertrandite, are found in Argentina, Brazil, India, Madagascar, Russia and the United States. Total world reserves of beryllium ore are greater than 400,000 tonnes.
Production
The extraction of beryllium from its compounds is a difficult process due to its high affinity for oxygen at elevated temperatures, and its ability to reduce water when its oxide film is removed. Currently the United States, China and Kazakhstan are the only three countries involved in the industrial-scale extraction of beryllium. Kazakhstan produces beryllium from a concentrate stockpiled before the breakup of the Soviet Union around 1991. This resource had become nearly depleted by mid-2010s.
Production of beryllium in Russia was halted in 1997, and is planned to be resumed in the 2020s.
Beryllium is most commonly extracted from the mineral beryl, which is either sintered using an extraction agent or melted into a soluble mixture. The sintering process involves mixing beryl with sodium fluorosilicate and soda at to form sodium fluoroberyllate, aluminium oxide and silicon dioxide. Beryllium hydroxide is precipitated from a solution of sodium fluoroberyllate and sodium hydroxide in water. The extraction of beryllium using the melt method involves grinding beryl into a powder and heating it to . The melt is quickly cooled with water and then reheated in concentrated sulfuric acid, mostly yielding beryllium sulfate and aluminium sulfate. Aqueous ammonia is then used to remove the aluminium and sulfur, leaving beryllium hydroxide. | Beryllium | Wikipedia | 446 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Beryllium hydroxide created using either the sinter or melt method is then converted into beryllium fluoride or beryllium chloride. To form the fluoride, aqueous ammonium hydrogen fluoride is added to beryllium hydroxide to yield a precipitate of ammonium tetrafluoroberyllate, which is heated to to form beryllium fluoride. Heating the fluoride to with magnesium forms finely divided beryllium, and additional heating to creates the compact metal. Heating beryllium hydroxide forms beryllium oxide, which becomes beryllium chloride when combined with carbon and chlorine. Electrolysis of molten beryllium chloride is then used to obtain the metal.
Chemical properties
Beryllium has a high electronegativity compared to other group 2 elements; thus C-Be bonds are less highly polarized than other C-MII bonds, although the attached carbon still bears a negative dipole moment.
A beryllium atom has the electronic configuration [He] 2s2. The predominant oxidation state of beryllium is +2; the beryllium atom has lost both of its valence electrons. Lower oxidation states complexes of beryllium are exceedingly rare. For example, bis(carbene) compounds proposed to contain beryllium in the 0 and +1 oxidation state have been reported, although these claims have proved controversial.
A stable complex with a Be-Be bond, which formally features beryllium in the +1 oxidation state, has been described. Beryllium's chemical behavior is largely a result of its small atomic and ionic radii. It thus has very high ionization potentials and strong polarization while bonded to other atoms, which is why all of its compounds are covalent. Its chemistry has similarities to that of aluminium, an example of a diagonal relationship. | Beryllium | Wikipedia | 386 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
At room temperature, the surface of beryllium forms a 1−10 nm-thick oxide passivation layer that prevents further reactions with air, except for gradual thickening of the oxide up to about 25 nm. When heated above about 500 °C, oxidation into the bulk metal progresses along grain boundaries. Once the metal is ignited in air by heating above the oxide melting point around 2500 °C, beryllium burns brilliantly, forming a mixture of beryllium oxide and beryllium nitride. Beryllium dissolves readily in non-oxidizing acids, such as HCl and diluted H2SO4, but not in nitric acid or water as this forms the oxide. This behavior is similar to that of aluminium. Beryllium also dissolves in alkali solutions.
Binary compounds of beryllium(II) are polymeric in the solid state. BeF2 has a silica-like structure with corner-shared BeF4 tetrahedra. BeCl2 and BeBr2 have chain structures with edge-shared tetrahedra. Beryllium oxide, BeO, is a white refractory solid which has a wurtzite crystal structure and a thermal conductivity as high as some metals. BeO is amphoteric. Beryllium sulfide, selenide and telluride are known, all having the zincblende structure. Beryllium nitride, Be3N2, is a high-melting-point compound which is readily hydrolyzed. Beryllium azide, BeN6 is known and beryllium phosphide, Be3P2 has a similar structure to Be3N2. A number of beryllium borides are known, such as Be5B, Be4B, Be2B, BeB2, BeB6 and BeB12. Beryllium carbide, Be2C, is a refractory brick-red compound that reacts with water to give methane. No beryllium silicide has been identified. | Beryllium | Wikipedia | 422 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
The halides BeX2 (X = F, Cl, Br, and I) have a linear monomeric molecular structure in the gas phase. Complexes of the halides are formed with one or more ligands donating a total of two pairs of electrons. Such compounds obey the octet rule. Other 4-coordinate complexes, such as the aqua-ion [Be(H2O)4]2+ also obey the octet rule.
Aqueous solutions
Solutions of beryllium salts, such as beryllium sulfate and beryllium nitrate, are acidic because of hydrolysis of the [Be(H2O)4]2+ ion. The concentration of the first hydrolysis product, [Be(H2O)3(OH)]+, is less than 1% of the beryllium concentration. The most stable hydrolysis product is the trimeric ion [Be3(OH)3(H2O)6]3+. Beryllium hydroxide, Be(OH)2, is insoluble in water at pH 5 or more. Consequently, beryllium compounds are generally insoluble at biological pH. Because of this, inhalation of beryllium metal dust leads to the development of the fatal condition of berylliosis. Be(OH)2 dissolves in strongly alkaline solutions.
Beryllium(II) forms few complexes with monodentate ligands because the water molecules in the aquo-ion, [Be(H2O)4]2+ are bound very strongly to the beryllium ion. Notable exceptions are the series of water-soluble complexes with the fluoride ion: [Be(H2O)4]^2+{} + \mathit{n}\,F^- <=> Be[(H2O)_{2\!-\mathit{n}}F_\mathit{n}]^{2\!-\mathit{n}}{} + \mathit{n}\,H2O | Beryllium | Wikipedia | 431 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Beryllium(II) forms many complexes with bidentate ligands containing oxygen-donor atoms. The species [Be3O(H2PO4)6]2- is notable for having a 3-coordinate oxide ion at its center. Basic beryllium acetate, Be4O(OAc)6, has an oxide ion surrounded by a tetrahedron of beryllium atoms.
With organic ligands, such as the malonate ion, the acid deprotonates when forming the complex. The donor atoms are two oxygens. H2A + [Be(H2O)4]^2+ <=> [BeA(H2O)2] + 2H+ + 2H2O H2A + [BeA(H2O)2] <=> [BeA2]^2- + 2H+ + 2H2O The formation of a complex is in competition with the metal ion-hydrolysis reaction and mixed complexes with both the anion and the hydroxide ion are also formed. For example, derivatives of the cyclic trimer are known, with a bidentate ligand replacing one or more pairs of water molecules.
Aliphatic hydroxycarboxylic acids such as glycolic acid form rather weak monodentate complexes in solution, in which the hydroxyl group remains intact. In the solid state, the hydroxyl group may deprotonate: a hexamer, Na_4[Be_6(OCH_2(O)O)_6] , was isolated long ago. Aromatic hydroxy ligands (i.e. phenols) form relatively strong complexes. For example, log K1 and log K2 values of 12.2 and 9.3 have been reported for complexes with tiron.
Beryllium has generally a rather poor affinity for ammine ligands. Ligands such as EDTA behave as dicarboxylic acids. There are many early reports of complexes with amino acids, but unfortunately they are not reliable as the concomitant hydrolysis reactions were not understood at the time of publication. Values for log β of ca. 6 to 7 have been reported. The degree of formation is small because of competition with hydrolysis reactions.
Organic chemistry | Beryllium | Wikipedia | 468 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Organoberyllium chemistry is limited to academic research due to the cost and toxicity of beryllium, beryllium derivatives and reagents required for the introduction of beryllium, such as beryllium chloride. Organometallic beryllium compounds are known to be highly reactive. Examples of known organoberyllium compounds are dineopentylberyllium, beryllocene (Cp2Be), diallylberyllium (by exchange reaction of diethyl beryllium with triallyl boron), bis(1,3-trimethylsilylallyl)beryllium, Be(mes)2, and (beryllium(I) complex) diberyllocene. Ligands can also be aryls and alkynyls.
History
The mineral beryl, which contains beryllium, has been used at least since the Ptolemaic dynasty of Egypt. In the first century CE, Roman naturalist Pliny the Elder mentioned in his encyclopedia Natural History that beryl and emerald ("smaragdus") were similar. The Papyrus Graecus Holmiensis, written in the third or fourth century CE, contains notes on how to prepare artificial emerald and beryl.
Early analyses of emeralds and beryls by Martin Heinrich Klaproth, Torbern Olof Bergman, Franz Karl Achard, and always yielded similar elements, leading to the mistaken conclusion that both substances are aluminium silicates. Mineralogist René Just Haüy discovered that both crystals are geometrically identical, and he asked chemist Louis-Nicolas Vauquelin for a chemical analysis.
In a 1798 paper read before the Institut de France, Vauquelin reported that he found a new "earth" by dissolving aluminium hydroxide from emerald and beryl in an additional alkali. The editors of the journal Annales de chimie et de physique named the new earth "glucine" for the sweet taste of some of its compounds. Klaproth preferred the name "beryllina" due to the fact that yttria also formed sweet salts. The name beryllium was first used by Friedrich Wöhler in 1828. | Beryllium | Wikipedia | 452 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Friedrich Wöhler and Antoine Bussy independently isolated beryllium in 1828 by the chemical reaction of metallic potassium with beryllium chloride, as follows:
BeCl2 + 2 K → 2 KCl + Be
Using an alcohol lamp, Wöhler heated alternating layers of beryllium chloride and potassium in a wired-shut platinum crucible. The above reaction immediately took place and caused the crucible to become white hot. Upon cooling and washing the resulting gray-black powder, he saw that it was made of fine particles with a dark metallic luster. The highly reactive potassium had been produced by the electrolysis of its compounds, a process discovered 21 years earlier. The chemical method using potassium yielded only small grains of beryllium from which no ingot of metal could be cast or hammered.
The direct electrolysis of a molten mixture of beryllium fluoride and sodium fluoride by Paul Lebeau in 1898 resulted in the first pure (99.5 to 99.8%) samples of beryllium. However, industrial production started only after the First World War. The original industrial involvement included subsidiaries and scientists related to the Union Carbide and Carbon Corporation in Cleveland, Ohio, and Siemens & Halske AG in Berlin. In the US, the process was ruled by Hugh S. Cooper, director of The Kemet Laboratories Company. In Germany, the first commercially successful process for producing beryllium was developed in 1921 by Alfred Stock and Hans Goldschmidt.
A sample of beryllium was bombarded with alpha rays from the decay of radium in a 1932 experiment by James Chadwick that uncovered the existence of the neutron. This same method is used in one class of radioisotope-based laboratory neutron sources that produce 30 neutrons for every million α particles.
Beryllium production saw a rapid increase during World War II due to the rising demand for hard beryllium-copper alloys and phosphors for fluorescent lights. Most early fluorescent lamps used zinc orthosilicate with varying content of beryllium to emit greenish light. Small additions of magnesium tungstate improved the blue part of the spectrum to yield an acceptable white light. Halophosphate-based phosphors replaced beryllium-based phosphors after beryllium was found to be toxic. | Beryllium | Wikipedia | 471 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Electrolysis of a mixture of beryllium fluoride and sodium fluoride was used to isolate beryllium during the 19th century. The metal's high melting point makes this process more energy-consuming than corresponding processes used for the alkali metals. Early in the 20th century, the production of beryllium by the thermal decomposition of beryllium iodide was investigated following the success of a similar process for the production of zirconium, but this process proved to be uneconomical for volume production.
Pure beryllium metal did not become readily available until 1957, even though it had been used as an alloying metal to harden and toughen copper much earlier. Beryllium could be produced by reducing beryllium compounds such as beryllium chloride with metallic potassium or sodium. Currently, most beryllium is produced by reducing beryllium fluoride with magnesium. The price on the American market for vacuum-cast beryllium ingots was about $338 per pound ($745 per kilogram) in 2001.
Between 1998 and 2008, the world's production of beryllium had decreased from 343 to about 200 tonnes. It then increased to 230 metric tons by 2018, of which 170 tonnes came from the United States.
Etymology
Beryllium was named for the semiprecious mineral beryl, from which it was first isolated. The name beryllium was introduced by Wöhler in 1828.
Although Humphry Davy failed to isolate it, he proposed the name glucium for the new metal, derived from the name glucina for the earth it was found in; altered forms of this name, glucinium or glucinum (symbol Gl) continued to be used into the 20th century. Both beryllium and glucinum were used concurrently until 1949, when the IUPAC adopted beryllium as the standard name of the element.
Applications
Radiation windows | Beryllium | Wikipedia | 396 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Because of its low atomic number and very low absorption for X-rays, the oldest and still one of the most important applications of beryllium is in radiation windows for X-ray tubes. Extreme demands are placed on purity and cleanliness of beryllium to avoid artifacts in the X-ray images. Thin beryllium foils are used as radiation windows for X-ray detectors, and their extremely low absorption minimizes the heating effects caused by high-intensity, low energy X-rays typical of synchrotron radiation. Vacuum-tight windows and beam-tubes for radiation experiments on synchrotrons are manufactured exclusively from beryllium. In scientific setups for various X-ray emission studies (e.g., energy-dispersive X-ray spectroscopy) the sample holder is usually made of beryllium because its emitted X-rays have much lower energies (≈100 eV) than X-rays from most studied materials.
Low atomic number also makes beryllium relatively transparent to energetic particles. Therefore, it is used to build the beam pipe around the collision region in particle physics setups, such as all four main detector experiments at the Large Hadron Collider (ALICE, ATLAS, CMS, LHCb), the Tevatron and at SLAC. The low density of beryllium allows collision products to reach the surrounding detectors without significant interaction, its stiffness allows a powerful vacuum to be produced within the pipe to minimize interaction with gases, its thermal stability allows it to function correctly at temperatures of only a few degrees above absolute zero, and its diamagnetic nature keeps it from interfering with the complex multipole magnet systems used to steer and focus the particle beams.
Mechanical applications
Because of its stiffness, light weight and dimensional stability over a wide temperature range, beryllium metal is used for lightweight structural components in the defense and aerospace industries in high-speed aircraft, guided missiles, spacecraft, and satellites, including the James Webb Space Telescope. Several liquid-fuel rockets have used rocket nozzles made of pure beryllium. Beryllium powder was itself studied as a rocket fuel, but this use has never materialized. A small number of extreme high-end bicycle frames have been built with beryllium. From 1998 to 2000, the McLaren Formula One team used Mercedes-Benz engines with beryllium-aluminium alloy pistons. The use of beryllium engine components was banned following a protest by Scuderia Ferrari. | Beryllium | Wikipedia | 502 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Mixing about 2.0% beryllium into copper forms an alloy called beryllium copper that is six times stronger than copper alone. Beryllium alloys are used in many applications because of their combination of elasticity, high electrical conductivity and thermal conductivity, high strength and hardness, nonmagnetic properties, as well as good corrosion and fatigue resistance. These applications include non-sparking tools that are used near flammable gases (beryllium nickel), springs, membranes (beryllium nickel and beryllium iron) used in surgical instruments, and high temperature devices. As little as 50 parts per million of beryllium alloyed with liquid magnesium leads to a significant increase in oxidation resistance and decrease in flammability.
The high elastic stiffness of beryllium has led to its extensive use in precision instrumentation, e.g. in inertial guidance systems and in the support mechanisms for optical systems. Beryllium-copper alloys were also applied as a hardening agent in "Jason pistols", which were used to strip the paint from the hulls of ships.
In sound amplification systems, the speed at which sound travels directly affects the resonant frequency of the amplifier, thereby influencing the range of audible high-frequency sounds. Beryllium stands out due to its exceptionally high speed of sound propagation compared to other metals. This unique property allows beryllium to achieve higher resonant frequencies, making it an ideal material for use as a diaphragm in high-quality loudspeakers.
Beryllium was used for cantilevers in high-performance phonograph cartridge styli, where its extreme stiffness and low density allowed for tracking weights to be reduced to 1 gram while still tracking high frequency passages with minimal distortion.
An earlier major application of beryllium was in brakes for military airplanes because of its hardness, high melting point, and exceptional ability to dissipate heat. Environmental considerations have led to substitution by other materials.
To reduce costs, beryllium can be alloyed with significant amounts of aluminium, resulting in the AlBeMet alloy (a trade name). This blend is cheaper than pure beryllium, while still retaining many desirable properties. | Beryllium | Wikipedia | 454 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Mirrors
Beryllium mirrors are of particular interest. Large-area mirrors, frequently with a honeycomb support structure, are used, for example, in meteorological satellites where low weight and long-term dimensional stability are critical. Smaller beryllium mirrors are used in optical guidance systems and in fire-control systems, e.g. in the German-made Leopard 1 and Leopard 2 main battle tanks. In these systems, very rapid movement of the mirror is required, which again dictates low mass and high rigidity. Usually the beryllium mirror is coated with hard electroless nickel plating which can be more easily polished to a finer optical finish than beryllium. In some applications, the beryllium blank is polished without any coating. This is particularly applicable to cryogenic operation where thermal expansion mismatch can cause the coating to buckle.
The James Webb Space Telescope has 18 hexagonal beryllium sections for its mirrors, each plated with a thin layer of gold. Because JWST will face a temperature of 33 K, the mirror is made of gold-plated beryllium, which is capable of handling extreme cold better than glass. Beryllium contracts and deforms less than glass and remains more uniform in such temperatures. For the same reason, the optics of the Spitzer Space Telescope are entirely built of beryllium metal.
Magnetic applications
Beryllium is non-magnetic. Therefore, tools fabricated out of beryllium-based materials are used by naval or military explosive ordnance disposal teams for work on or near naval mines, since these mines commonly have magnetic fuzes. They are also found in maintenance and construction materials near magnetic resonance imaging (MRI) machines because of the high magnetic fields generated. In the fields of radio communications and powerful (usually military) radars, hand tools made of beryllium are used to tune the highly magnetic klystrons, magnetrons, traveling wave tubes, etc., that are used for generating high levels of microwave power in the transmitters.
Nuclear applications
Thin plates or foils of beryllium are sometimes used in nuclear weapon designs as the very outer layer of the plutonium pits in the primary stages of thermonuclear bombs, placed to surround the fissile material. These layers of beryllium are good "pushers" for the implosion of the plutonium-239, and they are good neutron reflectors, just as in beryllium-moderated nuclear reactors. | Beryllium | Wikipedia | 503 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Beryllium is commonly used in some neutron sources in laboratory devices in which relatively few neutrons are needed (rather than having to use a nuclear reactor or a particle accelerator-powered neutron generator). For this purpose, a target of beryllium-9 is bombarded with energetic alpha particles from a radioisotope such as polonium-210, radium-226, plutonium-238, or americium-241. In the nuclear reaction that occurs, a beryllium nucleus is transmuted into carbon-12, and one free neutron is emitted, traveling in about the same direction as the alpha particle was heading. Such alpha decay-driven beryllium neutron sources, named "urchin" neutron initiators, were used in some early atomic bombs. Neutron sources in which beryllium is bombarded with gamma rays from a gamma decay radioisotope, are also used to produce laboratory neutrons.
Beryllium is used in fuel fabrication for CANDU reactors. The fuel elements have small appendages that are resistance brazed to the fuel cladding using an induction brazing process with Be as the braze filler material. Bearing pads are brazed in place to prevent contact between the fuel bundle and the pressure tube containing it, and inter-element spacer pads are brazed on to prevent element to element contact.
Beryllium is used at the Joint European Torus nuclear-fusion research laboratory, and it will be used in the more advanced ITER to condition the components which face the plasma. Beryllium has been proposed as a cladding material for nuclear fuel rods, because of its good combination of mechanical, chemical, and nuclear properties. Beryllium fluoride is one of the constituent salts of the eutectic salt mixture FLiBe, which is used as a solvent, moderator and coolant in many hypothetical molten salt reactor designs, including the liquid fluoride thorium reactor (LFTR).
Acoustics
The low weight and high rigidity of beryllium make it useful as a material for high-frequency speaker drivers. Because beryllium is expensive (many times more than titanium), hard to shape due to its brittleness, and toxic if mishandled, beryllium tweeters are limited to high-end home, pro audio, and public address applications. Some high-fidelity products have been fraudulently claimed to be made of the material. | Beryllium | Wikipedia | 494 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Some high-end phonograph cartridges used beryllium cantilevers to improve tracking by reducing mass.
Electronic
Beryllium is a p-type dopant in III-V compound semiconductors. It is widely used in materials such as GaAs, AlGaAs, InGaAs and InAlAs grown by molecular beam epitaxy (MBE). Cross-rolled beryllium sheet is an excellent structural support for printed circuit boards in surface-mount technology. In critical electronic applications, beryllium is both a structural support and heat sink. The application also requires a coefficient of thermal expansion that is well matched to the alumina and polyimide-glass substrates. The beryllium-beryllium oxide composite "E-Materials" have been specially designed for these electronic applications and have the additional advantage that the thermal expansion coefficient can be tailored to match diverse substrate materials.
Beryllium oxide is useful for many applications that require the combined properties of an electrical insulator and an excellent heat conductor, with high strength and hardness and a very high melting point. Beryllium oxide is frequently used as an insulator base plate in high-power transistors in radio frequency transmitters for telecommunications. Beryllium oxide is being studied for use in increasing the thermal conductivity of uranium dioxide nuclear fuel pellets. Beryllium compounds were used in fluorescent lighting tubes, but this use was discontinued because of the disease berylliosis which developed in the workers who were making the tubes.
Medical applications
Beryllium is a component of several dental alloys. Beryllium is used in X-ray windows because it is transparent to X-rays, allowing for clearer and more efficient imaging. In medical imaging equipment, such as CT scanners and mammography machines, beryllium's strength and light weight enhance durability and performance. Beryllium is used in analytical equipment for blood, HIV, and other diseases. Beryllium alloys are used in surgical instruments, optical mirrors, and laser systems for medical treatments.
Toxicity and safety | Beryllium | Wikipedia | 416 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Biological effects
Approximately 35 micrograms of beryllium is found in the average human body, an amount not considered harmful. Beryllium is chemically similar to magnesium and therefore can displace it from enzymes, which causes them to malfunction. Because Be2+ is a highly charged and small ion, it can easily get into many tissues and cells, where it specifically targets cell nuclei, inhibiting many enzymes, including those used for synthesizing DNA. Its toxicity is exacerbated by the fact that the body has no means to control beryllium levels, and once inside the body, beryllium cannot be removed.
Inhalation
Chronic beryllium disease (CBD), or berylliosis, is a pulmonary and systemic granulomatous disease caused by inhalation of dust or fumes contaminated with beryllium; either large amounts over a short time or small amounts over a long time can lead to this ailment. Symptoms of the disease can take up to five years to develop; about a third of patients with it die and the survivors are left disabled. The International Agency for Research on Cancer (IARC) lists beryllium and beryllium compounds as Category 1 carcinogens.
Occupational exposure
In the US, the Occupational Safety and Health Administration (OSHA) has designated a permissible exposure limit (PEL) for beryllium and beryllium compounds of 0.2 μg/m3 as an 8-hour time-weighted average (TWA) and 2.0 μg/m3 as a short-term exposure limit over a sampling period of 15 minutes. The National Institute for Occupational Safety and Health (NIOSH) has set a recommended exposure limit (REL) upper-bound threshold of 0.5 μg/m3. The IDLH (immediately dangerous to life and health) value is 4 mg/m3. The toxicity of beryllium is on par with other toxic metalloids/metals, such as arsenic and mercury.
Exposure to beryllium in the workplace can lead to a sensitized immune response, and over time development of berylliosis. NIOSH in the United States researches these effects in collaboration with a major manufacturer of beryllium products. NIOSH also conducts genetic research on sensitization and CBD, independently of this collaboration. | Beryllium | Wikipedia | 479 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
Acute beryllium disease in the form of chemical pneumonitis was first reported in Europe in 1933 and in the United States in 1943. A survey found that about 5% of workers in plants manufacturing fluorescent lamps in 1949 in the United States had beryllium-related lung diseases. Chronic berylliosis resembles sarcoidosis in many respects, and the differential diagnosis is often difficult. It killed some early workers in nuclear weapons design, such as Herbert L. Anderson.
Beryllium may be found in coal slag. When the slag is formulated into an abrasive agent for blasting paint and rust from hard surfaces, the beryllium can become airborne and become a source of exposure.
Although the use of beryllium compounds in fluorescent lighting tubes was discontinued in 1949, potential for exposure to beryllium exists in the nuclear and aerospace industries, in the refining of beryllium metal and the melting of beryllium-containing alloys, in the manufacturing of electronic devices, and in the handling of other beryllium-containing material.
Detection
Early researchers undertook the highly hazardous practice of identifying beryllium and its various compounds from its sweet taste. A modern test for beryllium in air and on surfaces has been developed and published as an international voluntary consensus standard, ASTM D7202. The procedure uses dilute ammonium bifluoride for dissolution and fluorescence detection with beryllium bound to sulfonated hydroxybenzoquinoline, allowing up to 100 times more sensitive detection than the recommended limit for beryllium concentration in the workplace. Fluorescence increases with increasing beryllium concentration. The new procedure has been successfully tested on a variety of surfaces and is effective for the dissolution and detection of refractory beryllium oxide and siliceous beryllium in minute concentrations (ASTM D7458). The NIOSH Manual of Analytical Methods contains methods for measuring occupational exposures to beryllium. | Beryllium | Wikipedia | 399 | 3378 | https://en.wikipedia.org/wiki/Beryllium | Physical sciences | Chemical elements_2 | null |
A bridge is a structure built to span a physical obstacle (such as a body of water, valley, road, or railway) without blocking the path underneath. It is constructed for the purpose of providing passage over the obstacle, which is usually something that is otherwise difficult or impossible to cross. There are many different designs of bridges, each serving a particular purpose and applicable to different situations. Designs of bridges vary depending on factors such as the function of the bridge, the nature of the terrain where the bridge is constructed and anchored, the material used to make it, and the funds available to build it.
The earliest bridges were likely made with fallen trees and stepping stones. The Neolithic people built boardwalk bridges across marshland. The Arkadiko Bridge, dating from the 13th century BC, in the Peloponnese is one of the oldest arch bridges in existence and use.
Etymology
The Oxford English Dictionary traces the origin of the word bridge to an Old English word brycg, of the same meaning.
The Oxford English Dictionary also notes that there is some suggestion that the word can be traced directly back to Proto-Indo-European *bʰrēw-. However, they also note that "this poses semantic problems."
The origin of the word for the card game of the same name is unknown, but may be from folk etymology.
History
The simplest and earliest types of bridges were stepping stones.
Neolithic people also built a form of boardwalk across marshes; examples of such bridges include the Sweet Track and the Post Track in England, approximately 6000 years old. Ancient people would also have used log bridges consisting of logs that fell naturally or were intentionally felled or placed across streams. Some of the first human-made bridges with significant span were probably intentionally felled trees. Among the oldest timber bridges is the Holzbrücke Rapperswil-Hurden bridge that crossed upper Lake Zürich in Switzerland; prehistoric timber pilings discovered to the west of the Seedamm causeway date back to 1523 BC. The first wooden footbridge there led across Lake Zürich; it was reconstructed several times through the late 2nd century AD, when the Roman Empire built a wooden bridge to carry transport across the lake. Between 1358 and 1360, Rudolf IV, Duke of Austria, built a 'new' wooden bridge across the lake that was used until 1878; it was approximately long and wide. On 6 April 2001, a reconstruction of the original wooden footbridge was opened; it is also the longest wooden bridge in Switzerland. | Bridge | Wikipedia | 500 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
The Arkadiko Bridge is one of four Mycenaean corbel arch bridges part of a former network of roads, designed to accommodate chariots, between the fort of Tiryns and town of Epidauros in the Peloponnese, in southern Greece. Dating to the Greek Bronze Age (13th century BC), it is one of the oldest arch bridges still in existence and use. Several intact, arched stone bridges from the Hellenistic era can be found in the Peloponnese.
The greatest bridge builders of antiquity were the ancient Romans. The Romans built arch bridges and aqueducts that could stand in conditions that would damage or destroy earlier designs, some of which still stand today. An example is the Alcántara Bridge, built over the river Tagus, in Spain. The Romans also used cement, which reduced the variation of strength found in natural stone. One type of cement, called pozzolana, consisted of water, lime, sand, and volcanic rock. Brick and mortar bridges were built after the Roman era, as the technology for cement was lost (then later rediscovered).
In India, the Arthashastra treatise by Kautilya mentions the construction of dams and bridges. A Mauryan bridge near Girnar was surveyed by James Princep. The bridge was swept away during a flood, and later repaired by Puspagupta, the chief architect of emperor Chandragupta I. The use of stronger bridges using plaited bamboo and iron chain was visible in India by about the 4th century. A number of bridges, both for military and commercial purposes, were constructed by the Mughal administration in India.
Although large bridges of wooden construction existed in China at the time of the Warring States period, the oldest surviving stone bridge in China is the Zhaozhou Bridge, built from 595 to 605 AD during the Sui dynasty. This bridge is also historically significant as it is the world's oldest open-spandrel stone segmental arch bridge. European segmental arch bridges date back to at least the Alconétar Bridge (approximately 2nd century AD), while the enormous Roman era Trajan's Bridge (105 AD) featured open-spandrel segmental arches in wooden construction.
Rope bridges, a simple type of suspension bridge, were used by the Inca civilization in the Andes mountains of South America, just prior to European colonization in the 16th century. | Bridge | Wikipedia | 488 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
The Ashanti built bridges over streams and rivers. They were constructed by pounding four large forked tree trunks into the stream bed, placing beams along these forked pillars, then positioning cross-beams that were finally covered with four to six inches of dirt.
During the 18th century, there were many innovations in the design of timber bridges by Hans Ulrich Grubenmann, Johannes Grubenmann, as well as others. The first book on bridge engineering was written by Hubert Gautier in 1716.
A major breakthrough in bridge technology came with the erection of the Iron Bridge in Shropshire, England in 1779. It used cast iron for the first time as arches to cross the river Severn. With the Industrial Revolution in the 19th century, truss systems of wrought iron were developed for larger bridges, but iron does not have the tensile strength to support large loads. With the advent of steel, which has a high tensile strength, much larger bridges were built, many using the ideas of Gustave Eiffel.
In Canada and the United States, numerous timber covered bridges were built in the late 1700s to the late 1800s, reminiscent of earlier designs in Germany and Switzerland. Some covered bridges were also built in Asia. In later years, some were partly made of stone or metal but the trusses were usually still made of wood; in the United States, there were three styles of trusses, the Queen Post, the Burr Arch and the Town Lattice. Hundreds of these structures still stand in North America. They were brought to the attention of the general public in the 1990s by the novel, movie and play The Bridges of Madison County.
In 1927, welding pioneer Stefan Bryła designed the first welded road bridge in the world, the Maurzyce Bridge which was later built across the river Słudwia at Maurzyce near Łowicz, Poland in 1929. In 1995, the American Welding Society presented the Historic Welded Structure Award for the bridge to Poland.
Types of bridges
Bridges can be categorized in several different ways. Common categories include the type of structural elements used, by what they carry, whether they are fixed or movable, and by the materials used. | Bridge | Wikipedia | 438 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Structure types
Bridges may be classified by how the actions of tension, compression, bending, torsion and shear are distributed through their structure. Most bridges will employ all of these to some degree, but only a few will predominate. The separation of forces and moments may be quite clear. In a suspension or cable-stayed bridge, the elements in tension are distinct in shape and placement. In other cases the forces may be distributed among a large number of members, as in a truss.
Some Engineers sub-divide 'beam' bridges into slab, beam-and-slab and box girder on the basis of their cross-section. A slab can be solid or voided (though this is no longer favored for inspectability reasons) while beam-and-slab consists of concrete or steel girders connected by a concrete slab. A box-girder cross-section consists of a single-cell or multi-cellular box. In recent years, integral bridge construction has also become popular.
Fixed or movable bridges
Most bridges are fixed bridges, meaning they have no moving parts and stay in one place until they fail or are demolished. Temporary bridges, such as Bailey bridges, are designed to be assembled, taken apart, transported to a different site, and re-used. They are important in military engineering and are also used to carry traffic while an old bridge is being rebuilt. Movable bridges are designed to move out of the way of boats or other kinds of traffic, which would otherwise be too tall to fit. These are generally electrically powered.
The Tank bridge transporter (TBT) has the same cross-country performance as a tank even when fully loaded. It can deploy, drop off and load bridges independently, but it cannot recover them.
Double-decked bridges | Bridge | Wikipedia | 358 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Double-decked (or double-decker) bridges have two levels, such as the George Washington Bridge, connecting New York City to Bergen County, New Jersey, US, as the world's busiest bridge, carrying 102 million vehicles annually; truss work between the roadway levels provided stiffness to the roadways and reduced movement of the upper level when the lower level was installed three decades after the upper level. The Tsing Ma Bridge and Kap Shui Mun Bridge in Hong Kong have six lanes on their upper decks, and on their lower decks there are two lanes and a pair of tracks for MTR metro trains. Some double-decked bridges only use one level for street traffic; the Washington Avenue Bridge in Minneapolis reserves its lower level for automobile and light rail traffic and its upper level for pedestrian and bicycle traffic (predominantly students at the University of Minnesota). Likewise, in Toronto, the Prince Edward Viaduct has five lanes of motor traffic, bicycle lanes, and sidewalks on its upper deck; and a pair of tracks for the Bloor–Danforth subway line on its lower deck. The western span of the San Francisco–Oakland Bay Bridge also has two levels.
Robert Stephenson's High Level Bridge across the River Tyne in Newcastle upon Tyne, completed in 1849, is an early example of a double-decked bridge. The upper level carries a railway, and the lower level is used for road traffic. Other examples include Britannia Bridge over the Menai Strait and Craigavon Bridge in Derry, Northern Ireland. The Oresund Bridge between Copenhagen and Malmö consists of a four-lane highway on the upper level and a pair of railway tracks at the lower level. Tower Bridge in London is different example of a double-decked bridge, with the central section consisting of a low-level bascule span and a high-level footbridge.
Viaducts
A viaduct is made up of multiple bridges connected into one longer structure. The longest and some of the highest bridges are viaducts, such as the Lake Pontchartrain Causeway and Millau Viaduct.
Multi-way bridge
A multi-way bridge has three or more separate spans which meet near the center of the bridge. Multi-way bridges with only three spans appear as a "T" or "Y" when viewed from above. Multi-way bridges are extremely rare. The Tridge, Margaret Bridge, and Zanesville Y-Bridge are examples. | Bridge | Wikipedia | 486 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Bridge types by use
A bridge can be categorized by what it is designed to carry, such as trains, pedestrian or road traffic (road bridge), a pipeline (Pipe bridge) or waterway for water transport or barge traffic. An aqueduct is a bridge that carries water, resembling a viaduct, which is a bridge that connects points of equal height. A road-rail bridge carries both road and rail traffic. Overway is a term for a bridge that separates incompatible intersecting traffic, especially road and rail.
Some bridges accommodate other purposes, such as the tower of Nový Most Bridge in Bratislava, which features a restaurant, or a bridge-restaurant which is a bridge built to serve as a restaurant. Other suspension bridge towers carry transmission antennas.
Conservationists use wildlife overpasses to reduce habitat fragmentation and animal-vehicle collisions. The first animal bridges sprung up in France in the 1950s, and these types of bridges are now used worldwide to protect both large and small wildlife.
Bridges are subject to unplanned uses as well. The areas underneath some bridges have become makeshift shelters and homes to homeless people, and the undertimbers of bridges all around the world are spots of prevalent graffiti. Some bridges attract people attempting suicide, and become known as suicide bridges.
Bridge types by material
The materials used to build the structure are also used to categorize bridges. Until the end of the 18th century, bridges were made out of timber, stone and masonry. Modern bridges are currently built in concrete, steel, fiber reinforced polymers (FRP), stainless steel or combinations of those materials. Living bridges have been constructed of live plants such as Ficus elastica tree roots in India and wisteria vines in Japan.
Analysis and design | Bridge | Wikipedia | 346 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Unlike buildings whose design is led by architects, bridges are usually designed by engineers. This follows from the importance of the engineering requirements; namely spanning the obstacle and having the durability to survive, with minimal maintenance, in an aggressive outdoor environment. Bridges are first analysed; the bending moment and shear force distributions are calculated due to the applied loads. For this, the finite element method is the most popular. The analysis can be one-, two-, or three-dimensional. For the majority of bridges, a two-dimensional plate model (often with stiffening beams) is sufficient or an upstand finite element model. On completion of the analysis, the bridge is designed to resist the applied bending moments and shear forces, section sizes are selected with sufficient capacity to resist the stresses. Many bridges are made of prestressed concrete which has good durability properties, either by pre-tensioning of beams prior to installation or post-tensioning on site.
In most countries, bridges, like other structures, are designed according to Load and Resistance Factor Design (LRFD) principles. In simple terms, this means that the load is factored up by a factor greater than unity, while the resistance or capacity of the structure is factored down, by a factor less than unity. The effect of the factored load (stress, bending moment) should be less than the factored resistance to that effect. Both of these factors allow for uncertainty and are greater when the uncertainty is greater.
Aesthetics
Most bridges are utilitarian in appearance, but in some cases, the appearance of the bridge can have great importance. Often, this is the case with a large bridge that serves as an entrance to a city, or crosses over a main harbor entrance. These are sometimes known as signature bridges. Designers of bridges in parks and along parkways often place more importance on aesthetics, as well. Examples include the stone-faced bridges along the Taconic State Parkway in New York. | Bridge | Wikipedia | 400 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Bridges are typically more aesthetically pleasing if they are simple in shape, the deck is thinner in proportion to its span, the lines of the structure are continuous, and the shapes of the structural elements reflect the forces acting on them. To create a beautiful image, some bridges are built much taller than necessary. This type, often found in east-Asian style gardens, is called a Moon bridge, evoking a rising full moon. Other garden bridges may cross only a dry bed of stream-washed pebbles, intended only to convey an impression of a stream. Often in palaces, a bridge will be built over an artificial waterway as symbolic of a passage to an important place or state of mind. A set of five bridges cross a sinuous waterway in an important courtyard of the Forbidden City in Beijing, China. The central bridge was reserved exclusively for the use of the Emperor and Empress, with their attendants.
Bridge maintenance
The estimated life of bridges varies between 25 and 80 years depending on location and material.
Bridges may age hundred years with proper maintenance and rehabilitation. Bridge maintenance consisting of a combination of structural health monitoring and testing. This is regulated in country-specific engineer standards and includes an ongoing monitoring every three to six months, a simple test or inspection every two to three years and a major inspection every six to ten years. In Europe, the cost of maintenance is considerable and is higher in some countries than spending on new bridges. The lifetime of welded steel bridges can be significantly extended by aftertreatment of the weld transitions. This results in a potential high benefit, using existing bridges far beyond the planned lifetime.
Bridge traffic loading
While the response of a bridge to the applied loading is well understood, the applied traffic loading itself is still the subject of research. This is a statistical problem as loading is highly variable, particularly for road bridges. Load Effects in bridges (stresses, bending moments) are designed for using the principles of Load and Resistance Factor Design. Before factoring to allow for uncertainty, the load effect is generally considered to be the maximum characteristic value in a specified return period. Notably, in Europe, it is the maximum value expected in 1000 years. | Bridge | Wikipedia | 432 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Bridge standards generally include a load model, deemed to represent the characteristic maximum load to be expected in the return period. In the past, these load models were agreed by standard drafting committees of experts but today, this situation is changing. It is now possible to measure the components of bridge traffic load, to weigh trucks, using weigh-in-motion (WIM) technologies. With extensive WIM databases, it is possible to calculate the maximum expected load effect in the specified return period. This is an active area of research, addressing issues of opposing direction lanes, side-by-side (same direction) lanes, traffic growth, permit/non-permit vehicles and long-span bridges (see below). Rather than repeat this complex process every time a bridge is to be designed, standards authorities specify simplified notional load models, notably HL-93, intended to give the same load effects as the characteristic maximum values. The Eurocode is an example of a standard for bridge traffic loading that was developed in this way.
Traffic loading on long span bridges
Most bridge standards are only applicable for short and medium spans - for example, the Eurocode is only applicable for loaded lengths up to 200 m. Longer spans are dealt with on a case-by-case basis. It is generally accepted that the intensity of load reduces as span increases because the probability of many trucks being closely spaced and extremely heavy reduces as the number of trucks involved increases. It is also generally assumed that short spans are governed by a small number of trucks traveling at high speed, with an allowance for dynamics. Longer spans on the other hand, are governed by congested traffic and no allowance for dynamics is needed.
Calculating the loading due to congested traffic remains a challenge as there is a paucity of data on inter-vehicle gaps, both within-lane and inter-lane, in congested conditions. Weigh-in-Motion (WIM) systems provide data on inter-vehicle gaps but only operate well in free flowing traffic conditions. Some authors have used cameras to measure gaps and vehicle lengths in jammed situations and have inferred weights from lengths using WIM data. Others have used microsimulation to generate typical clusters of vehicles on the bridge. | Bridge | Wikipedia | 446 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Bridge vibration
Bridges vibrate under load and this contributes, to a greater or lesser extent, to the stresses. Vibration and dynamics are generally more significant for slender structures such as pedestrian bridges and long-span road or rail bridges. One of the most famous examples is the Tacoma Narrows Bridge that collapsed shortly after being constructed due to excessive vibration. More recently, the Millennium Bridge in London vibrated excessively under pedestrian loading and was closed and retrofitted with a system of dampers. For smaller bridges, dynamics is not catastrophic but can contribute an added amplification to the stresses due to static effects. For example, the Eurocode for bridge loading specifies amplifications of between 10% and 70%, depending on the span, the number of traffic lanes and the type of stress (bending moment or shear force).
Vehicle-bridge dynamic interaction
There have been many studies of the dynamic interaction between vehicles and bridges during vehicle crossing events. Fryba did pioneering work on the interaction of a moving load and an Euler-Bernoulli beam. With increased computing power, vehicle-bridge interaction (VBI) models have become ever more sophisticated. The concern is that one of the many natural frequencies associated with the vehicle will resonate with the bridge's first natural frequency. The vehicle-related frequencies include body bounce and axle hop but there are also pseudo-frequencies associated with the vehicle's speed of crossing and there are many frequencies associated with the surface profile. Given the wide variety of heavy vehicles on road bridges, a statistical approach has been suggested, with VBI analyses carried out for many statically extreme loading events.
Bridge failures
The failure of bridges is of special concern for structural engineers in trying to learn lessons vital to bridge design, construction and maintenance.
The failure of bridges first assumed national interest in Britain during the Victorian era when many new designs were being built, often using new materials, with some of them failing catastrophically.
In the United States, the National Bridge Inventory tracks the structural evaluations of all bridges, including designations such as "structurally deficient" and "functionally obsolete". | Bridge | Wikipedia | 421 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Bridge health monitoring
There are several methods used to monitor the condition of large structures, like bridges. Many long-span bridges are now routinely monitored with a range of sensors, including strain transducers, accelerometers, tiltmeters, and GPS. Accelerometers have the advantage that they are inertial, i.e., they do not require a reference point to measure from. This is often a problem for distance or deflection measurement, especially if the bridge is over water. Crowdsourcing bridge conditions by accessing data passively captured by cell phones, which routinely include accelerometers and GPS sensors, has been suggested as an alternative to including sensors during bridge construction and an augment for professional examinations.
An option for structural-integrity monitoring is "non-contact monitoring", which uses the Doppler effect (Doppler shift). A laser beam from a Laser Doppler Vibrometer is directed at the point of interest, and the vibration amplitude and frequency are extracted from the Doppler shift of the laser beam frequency due to the motion of the surface. The advantage of this method is that the setup time for the equipment is faster and, unlike an accelerometer, this makes measurements possible on multiple structures in as short a time as possible. Additionally, this method can measure specific points on a bridge that might be difficult to access. However, vibrometers are relatively expensive and have the disadvantage that a reference point is needed to measure from.
Snapshots in time of the external condition of a bridge can be recorded using Lidar to aid bridge inspection. This can provide measurement of the bridge geometry (to facilitate the building of a computer model) but the accuracy is generally insufficient to measure bridge deflections under load.
While larger modern bridges are routinely monitored electronically, smaller bridges are generally inspected visually by trained inspectors. There is considerable research interest in the challenge of smaller bridges as they are often remote and do not have electrical power on site. Possible solutions are the installation of sensors on a specialist inspection vehicle and the use of its measurements as it drives over the bridge to infer information about the bridge condition. These vehicles can be equipped with accelerometers, gyrometers, Laser Doppler Vibrometers and some even have the capability to apply a resonant force to the road surface to dynamically excite the bridge at its resonant frequency.
Visual index | Bridge | Wikipedia | 501 | 3397 | https://en.wikipedia.org/wiki/Bridge | Technology | Transportation | null |
Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca (or Cladocera), Notostraca, the Devonian Lepidocaris and possibly the Cambrian Rehbachiella. They are mostly small, freshwater animals that feed on plankton and detritus.
Description
Members of the Branchiopoda are unified by the presence of gills on many of the animals' appendages, including some of the mouthparts. This is also responsible for the name of the group (from the , gills, akin to , windpipe; , foot). They generally possess compound eyes and a carapace, which may be a shell of two valves enclosing the trunk (as in most Cladocera), broad and shallow (as in the Notostraca), or entirely absent (as in the Anostraca). In the groups where the carapace prevents the use of the trunk limbs for swimming (Cladocera and clam shrimp), the antennae are used for locomotion, as they are in the nauplius. Male fairy shrimp have an enlarged pair of antennae with which they grasp the female during mating, while the bottom-feeding Notostraca, the antennae are reduced to vestiges. The trunk limbs are beaten in a metachronal rhythm, causing a flow of water along the midline of the animal, from which it derives oxygen, food and, in the case of the Anostraca and Notostraca, movement.
Ecology
Branchiopods are found in continental fresh water, including temporary pools and in hypersaline lakes, and some in brackish water. Only two groups of water fleas include marine species: Family Podonidae in the order Diplostraca, and family Sididae in the order Diplostraca.
Most branchiopodans eat floating detritus or plankton, which they take using the setae on their appendages. But notostracans are omnivorous and very opportunistic feeders and will eat algae and bacteria in addition to animals as both predators and scavengers.
Taxonomy
In early taxonomic treatments, the current members of the Branchiopoda were all placed in a single genus, Monoculus. The taxon Branchiopoda was erected by Pierre André Latreille in 1817, initially at the rank of order. | Branchiopoda | Wikipedia | 496 | 3406 | https://en.wikipedia.org/wiki/Branchiopoda | Biology and health sciences | Crustaceans | Animals |
The current upper-level classification of Branchiopoda, according to the World Register of Marine Species (2021), is as follows:
Class Branchiopoda Latreille, 1817
Subclass Sarsostraca Tasch, 1969
Order Anostraca Sars, 1867
Suborder Anostracina Weekers et al., 2002
Suborder Artemiina Weekers et al., 2002
Subclass Phyllopoda Preuss, 1951
Superorder Diplostraca Gerstaecker, 1866
Order Anomopoda G.O. Sars, 1865
Order Ctenopoda G.O. Sars, 1865
Order Cyclestherida Sars G.O., 1899
Order Haplopoda G.O. Sars, 1865
Order Laevicaudata Linder, 1945
Order Onychopoda G.O. Sars, 1865
Order Spinicaudata Linder, 1945
Order Notostraca G. O. Sars, 1867
Genus †Rehbachiella? Müller, 1983
In addition, the extinct genus Lepidocaris is generally placed in Branchiopoda.
Anostraca
The fairy shrimp of the order Anostraca are usually long (exceptionally up to ). Most species have 20 body segments, bearing 11 pairs of leaf-like phyllopodia (swimming legs), and the body lacks a carapace. They live in vernal pools and hypersaline lakes across the world, including pools in deserts, in ice-covered mountain lakes and in Antarctica. They swim "upside-down" and feed by filtering organic particles from the water or by scraping algae from surfaces. They are an important food for many birds and fish, and are cultured and harvested for use as fish food. There are 300 species spread across 8 families.
Lipostraca
Lipostraca contains a single extinct Early Devonian species, Lepidocaris rhyniensis, which is the most abundant animal in the Rhynie chert deposits. It resembles modern Anostraca, to which it is probably closely related, although its relationships to other orders remain unclear. The body is long, with 23 body segments and 19 pairs of appendages, but no carapace. It occurred chiefly among charophytes, probably in alkaline temporary pools.
Notostraca | Branchiopoda | Wikipedia | 483 | 3406 | https://en.wikipedia.org/wiki/Branchiopoda | Biology and health sciences | Crustaceans | Animals |
The order Notostraca comprises the single family Triopsidae, containing the tadpole shrimp or shield shrimp. The two genera, Triops and Lepidurus, are considered living fossils, having not changed significantly in outward form since the Triassic. They have a broad, flat carapace, which conceals the head and bears a single pair of compound eyes. The abdomen is long, appears to be segmented and bears numerous pairs of flattened legs. The telson is flanked by a pair of long, thin caudal rami. Phenotypic plasticity within taxa makes species-level identification difficult, and is further compounded by variation in the mode of reproduction. The evidence of phenotypic plasticity of Arctic tadpole shrimp (Lepidurus arcticus, Notostraca) has been observed in Svalbard. Notostracans are the largest branchiopodans and are omnivores living on the bottom of temporary pools, ponds and shallow lakes.
Laevicaudata, Spinicaudata and Cyclestherida (once Conchostraca)
Clam shrimp are bivalved animals which have lived since at least the Devonian. The three groups are not believed to form a clade. They have 10–32 trunk segments, decreasing in size from front to back, and each bears a pair of legs which also carry gills. A strong muscle can close the two halves of the shell together.
Anomopoda, Ctenopoda, Onychopoda, and Haplopoda (once Cladocera)
These four orders make up a group of small crustaceans commonly called water fleas. Around 620 species have been recognised so far, with many more undescribed. They are ubiquitous in inland aquatic habitats, but rare in the oceans. Most are long, with a down-turned head, and a carapace covering the apparently unsegmented thorax and abdomen. There is a single median compound eye. Most species show cyclical parthenogenesis, where asexual reproduction is occasionally supplemented by sexual reproduction, which produces resting eggs that allow the species to survive harsh conditions and disperse to distant habitats. In the water bodies of the world, a lot of Cladocera are non-native species, many of which pose a great threat to aquatic ecosystems. | Branchiopoda | Wikipedia | 471 | 3406 | https://en.wikipedia.org/wiki/Branchiopoda | Biology and health sciences | Crustaceans | Animals |
Evolution
The fossil record of branchiopods extends back at least into the Upper Cambrian and possibly further. The group is thought to be monophyletic, with the Anostraca having been the first group to branch off. It is thought that the group evolved in the seas, but was forced into temporary pools and hypersaline lakes by the evolution of bony fishes. Although they were previously considered the sister group to the remaining crustaceans, it is now widely accepted that crustaceans form a paraphyletic group, and Branchiopoda are thought to be sister to a clade comprising Xenocarida (Remipedia and Cephalocarida) and Hexapoda (insects and their relatives). | Branchiopoda | Wikipedia | 149 | 3406 | https://en.wikipedia.org/wiki/Branchiopoda | Biology and health sciences | Crustaceans | Animals |
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the bee hummingbird to the common ostrich. There are over 11,000 living species and they are split into 44 orders. More than half are passerine or "perching" birds. Birds have wings whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which are modified forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites, penguins, and diverse endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming. The study of birds is called ornithology.
Birds are feathered theropod dinosaurs and constitute the only known living dinosaurs. Likewise, birds are considered reptiles in the modern cladistic sense of the term, and their closest living relatives are the crocodilians. Birds are descendants of the primitive avialans (whose members include Archaeopteryx) which first appeared during the Late Jurassic. According to some estimates, modern birds (Neornithes) evolved in the Late Cretaceous or between the Early and Late Cretaceous (100 Ma) and diversified dramatically around the time of the Cretaceous–Paleogene extinction event 66 million years ago, which killed off the pterosaurs and all non-ornithuran dinosaurs. | Bird | Wikipedia | 360 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Many social species preserve knowledge across generations (culture). Birds are social, communicating with visual signals, calls, and songs, and participating in such behaviour as cooperative breeding and hunting, flocking, and mobbing of predators. The vast majority of bird species are socially (but not necessarily sexually) monogamous, usually for one breeding season at a time, sometimes for years, and rarely for life. Other species have breeding systems that are polygynous (one male with many females) or, rarely, polyandrous (one female with many males). Birds produce offspring by laying eggs which are fertilised through sexual reproduction. They are usually laid in a nest and incubated by the parents. Most birds have an extended period of parental care after hatching.
Many species of birds are economically important as food for human consumption and raw material in manufacturing, with domesticated and undomesticated birds being important sources of eggs, meat, and feathers. Songbirds, parrots, and other species are popular as pets. Guano (bird excrement) is harvested for use as a fertiliser. Birds figure throughout human culture. About 120 to 130 species have become extinct due to human activity since the 17th century, and hundreds more before then. Human activity threatens about 1,200 bird species with extinction, though efforts are underway to protect them. Recreational birdwatching is an important part of the ecotourism industry.
Evolution and classification
The first classification of birds was developed by Francis Willughby and John Ray in their 1676 volume Ornithologiae.
Carl Linnaeus modified that work in 1758 to devise the taxonomic classification system currently in use. Birds are categorised as the biological class Aves in Linnaean taxonomy. Phylogenetic taxonomy places Aves in the clade Theropoda as an infraclass or more recently a subclass. | Bird | Wikipedia | 381 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Definition
Aves and a sister group, the order Crocodilia, contain the only living representatives of the reptile clade Archosauria. During the late 1990s, Aves was most commonly defined phylogenetically as all descendants of the most recent common ancestor of modern birds and Archaeopteryx lithographica. However, an earlier definition proposed by Jacques Gauthier gained wide currency in the 21st century, and is used by many scientists including adherents to the PhyloCode. Gauthier defined Aves to include only the crown group of the set of modern birds. This was done by excluding most groups known only from fossils, and assigning them, instead, to the broader group Avialae, on the principle that a clade based on extant species should be limited to those extant species and their closest extinct relatives.
Gauthier and de Queiroz identified four different definitions for the same biological name "Aves", which is a problem. The authors proposed to reserve the term Aves only for the crown group consisting of the last common ancestor of all living birds and all of its descendants, which corresponds to meaning number 4 below. They assigned other names to the other groups.
Aves can mean all archosaurs closer to birds than to crocodiles (alternately Avemetatarsalia)
Aves can mean those advanced archosaurs with feathers (alternately Avifilopluma)
Aves can mean those feathered dinosaurs that fly (alternately Avialae)
Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "crown group", in this sense synonymous with Neornithes)
Under the fourth definition Archaeopteryx, traditionally considered one of the earliest members of Aves, is removed from this group, becoming a non-avian dinosaur instead. These proposals have been adopted by many researchers in the field of palaeontology and bird evolution, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is often used synonymously with the vernacular term "bird" by these researchers. | Bird | Wikipedia | 433 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Most researchers define Avialae as branch-based clade, though definitions vary. Many authors have used a definition similar to "all theropods closer to birds than to Deinonychus", with Troodon being sometimes added as a second external specifier in case it is closer to birds than to Deinonychus. Avialae is also occasionally defined as an apomorphy-based clade (that is, one based on physical characteristics). Jacques Gauthier, who named Avialae in 1986, re-defined it in 2001 as all dinosaurs that possessed feathered wings used in flapping flight, and the birds that descended from them.
Despite being currently one of the most widely used, the crown-group definition of Aves has been criticised by some researchers. Lee and Spencer (1997) argued that, contrary to what Gauthier defended, this definition would not increase the stability of the clade and the exact content of Aves will always be uncertain because any defined clade (either crown or not) will have few synapomorphies distinguishing it from its closest relatives. Their alternative definition is synonymous to Avifilopluma.
Dinosaurs and the origin of birds
Based on fossil and biological evidence, most scientists accept that birds are a specialised subgroup of theropod dinosaurs and, more specifically, members of Maniraptora, a group of theropods which includes dromaeosaurids and oviraptorosaurs, among others. As scientists have discovered more theropods closely related to birds, the previously clear distinction between non-birds and birds has become blurred. By the 2000s, discoveries in the Liaoning Province of northeast China, which demonstrated many small theropod feathered dinosaurs, contributed to this ambiguity. | Bird | Wikipedia | 357 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The consensus view in contemporary palaeontology is that the flying theropods, or avialans, are the closest relatives of the deinonychosaurs, which include dromaeosaurids and troodontids. Together, these form a group called Paraves. Some basal members of Deinonychosauria, such as Microraptor, have features which may have enabled them to glide or fly. The most basal deinonychosaurs were very small. This evidence raises the possibility that the ancestor of all paravians may have been arboreal, have been able to glide, or both. Unlike Archaeopteryx and the non-avialan feathered dinosaurs, who primarily ate meat, studies suggest that the first avialans were omnivores.
The Late Jurassic Archaeopteryx is well known as one of the first transitional fossils to be found, and it provided support for the theory of evolution in the late 19th century. Archaeopteryx was the first fossil to display both clearly traditional reptilian characteristics—teeth, clawed fingers, and a long, lizard-like tail—as well as wings with flight feathers similar to those of modern birds. It is not considered a direct ancestor of birds, though it is possibly closely related to the true ancestor.
Early evolution
Over 40% of key traits found in modern birds evolved during the 60 million year transition from the earliest bird-line archosaurs to the first maniraptoromorphs, i.e. the first dinosaurs closer to living birds than to Tyrannosaurus rex. The loss of osteoderms otherwise common in archosaurs and acquisition of primitive feathers might have occurred early during this phase. After the appearance of Maniraptoromorpha, the next 40 million years marked a continuous reduction of body size and the accumulation of neotenic (juvenile-like) characteristics. Hypercarnivory became increasingly less common while braincases enlarged and forelimbs became longer. The integument evolved into complex, pennaceous feathers.
The oldest known paravian (and probably the earliest avialan) fossils come from the Tiaojishan Formation of China, which has been dated to the late Jurassic period (Oxfordian stage), about 160 million years ago. The avialan species from this time period include Anchiornis huxleyi, Xiaotingia zhengi, and Aurornis xui. | Bird | Wikipedia | 502 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The well-known probable early avialan, Archaeopteryx, dates from slightly later Jurassic rocks (about 155 million years old) from Germany. Many of these early avialans shared unusual anatomical features that may be ancestral to modern birds but were later lost during bird evolution. These features include enlarged claws on the second toe which may have been held clear of the ground in life, and long feathers or "hind wings" covering the hind limbs and feet, which may have been used in aerial maneuvering.
Avialans diversified into a wide variety of forms during the Cretaceous period. Many groups retained primitive characteristics, such as clawed wings and teeth, though the latter were lost independently in a number of avialan groups, including modern birds (Aves). Increasingly stiff tails (especially the outermost half) can be seen in the evolution of maniraptoromorphs, and this process culminated in the appearance of the pygostyle, an ossification of fused tail vertebrae. In the late Cretaceous, about 100 million years ago, the ancestors of all modern birds evolved a more open pelvis, allowing them to lay larger eggs compared to body size. Around 95 million years ago, they evolved a better sense of smell.
A third stage of bird evolution starting with Ornithothoraces (the "bird-chested" avialans) can be associated with the refining of aerodynamics and flight capabilities, and the loss or co-ossification of several skeletal features. Particularly significant are the development of an enlarged, keeled sternum and the alula, and the loss of grasping hands.
Early diversity of bird ancestors
The first large, diverse lineage of short-tailed avialans to evolve were the Enantiornithes, or "opposite birds", so named because the construction of their shoulder bones was in reverse to that of modern birds. Enantiornithes occupied a wide array of ecological niches, from sand-probing shorebirds and fish-eaters to tree-dwelling forms and seed-eaters. While they were the dominant group of avialans during the Cretaceous period, Enantiornithes became extinct along with many other dinosaur groups at the end of the Mesozoic era. | Bird | Wikipedia | 457 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Many species of the second major avialan lineage to diversify, the Euornithes (meaning "true birds", because they include the ancestors of modern birds), were semi-aquatic and specialised in eating fish and other small aquatic organisms. Unlike the Enantiornithes, which dominated land-based and arboreal habitats, most early euornithians lacked perching adaptations and likely included shorebird-like species, waders, and swimming and diving species.
The latter included the superficially gull-like Ichthyornis and the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic. The early euornithians also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithians retained teeth in other parts of the jaws). Euornithes also included the first avialans to develop true pygostyle and a fully mobile fan of tail feathers, which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight.
A study on mosaic evolution in the avian skull found that the last common ancestor of all Neornithes might have had a beak similar to that of the modern hook-billed vanga and a skull similar to that of the Eurasian golden oriole. As both species are small aerial and canopy foraging omnivores, a similar ecological niche was inferred for this hypothetical ancestor.
Diversification of modern birds
Most studies agree on a Cretaceous age for the most recent common ancestor of modern birds but estimates range from the Early Cretaceous to the latest Cretaceous. Similarly, there is no agreement on whether most of the early diversification of modern birds occurred in the Cretaceous and associated with breakup of the supercontinent Gondwana or occurred later and potentially as a consequence of the Cretaceous–Palaeogene extinction event. This disagreement is in part caused by a divergence in the evidence; most molecular dating studies suggests a Cretaceous evolutionary radiation, while fossil evidence points to a Cenozoic radiation (the so-called 'rocks' versus 'clocks' controversy). | Bird | Wikipedia | 463 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The discovery in 2005 of Vegavis from the Maastrichtian, the last stage of the Late Cretaceous, proved that the diversification of modern birds started before the Cenozoic era. The affinities of an earlier fossil, the possible galliform Austinornis lentus, dated to about 85 million years ago, are still too controversial to provide a fossil evidence of modern bird diversification. In 2020, Asteriornis from the Maastrichtian was described, it appears to be a close relative of Galloanserae, the earliest diverging lineage within Neognathae.
Attempts to reconcile molecular and fossil evidence using genomic-scale DNA data and comprehensive fossil information have not resolved the controversy. However, a 2015 estimate that used a new method for calibrating molecular clocks confirmed that while modern birds originated early in the Late Cretaceous, likely in Western Gondwana, a pulse of diversification in all major groups occurred around the Cretaceous–Palaeogene extinction event. Modern birds would have expanded from West Gondwana through two routes. One route was an Antarctic interchange in the Paleogene. The other route was probably via Paleocene land bridges between South America and North America, which allowed for the rapid expansion and diversification of Neornithes into the Holarctic and Paleotropics. On the other hand, the occurrence of Asteriornis in the Northern Hemisphere suggest that Neornithes dispersed out of East Gondwana before the Paleocene.
Classification of bird orders
All modern birds lie within the crown group Aves (alternately Neornithes), which has two subdivisions: the Palaeognathae, which includes the flightless ratites (such as the ostriches) and the weak-flying tinamous, and the extremely diverse Neognathae, containing all other birds. These two subdivisions have variously been given the rank of superorder, cohort, or infraclass. The number of known living bird species is around 11,000 although sources may differ in their precise numbers.
Cladogram of modern bird relationships based on Stiller et al (2024)., showing the 44 orders recognised by the IOC. | Bird | Wikipedia | 445 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The classification of birds is a contentious issue. Sibley and Ahlquist's Phylogeny and Classification of Birds (1990) is a landmark work on the subject. Most evidence seems to suggest the assignment of orders is accurate, but scientists disagree about the relationships among the orders themselves; evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem, but no strong consensus has emerged. Fossil and molecular evidence from the 2010s is providing an increasingly clear picture of the evolution of modern bird orders.
Genomics
In 2010, the genome had been sequenced for only two birds, the chicken and the zebra finch. , the genomes of 542 species of birds had been completed. At least one genome has been sequenced from every order. These include at least one species in about 90% of extant avian families (218 out of 236 families recognised by the Howard and Moore Checklist).
Being able to sequence and compare whole genomes gives researchers many types of information, about genes, the DNA that regulates the genes, and their evolutionary history. This has led to reconsideration of some of the classifications that were based solely on the identification of protein-coding genes. Waterbirds such as pelicans and flamingos, for example, may have in common specific adaptations suited to their environment that were developed independently.
Distribution
Birds live and breed in most terrestrial habitats and on all seven continents, reaching their southern extreme in the snow petrel's breeding colonies up to inland in Antarctica. The highest bird diversity occurs in tropical regions. It was earlier thought that this high diversity was the result of higher speciation rates in the tropics; however studies from the 2000s found higher speciation rates in the high latitudes that were offset by greater extinction rates than in the tropics. Many species migrate annually over great distances and across oceans; several families of birds have adapted to life both on the world's oceans and in them, and some seabird species come ashore only to breed, while some penguins have been recorded diving up to deep. | Bird | Wikipedia | 417 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Many bird species have established breeding populations in areas to which they have been introduced by humans. Some of these introductions have been deliberate; the ring-necked pheasant, for example, has been introduced around the world as a game bird. Others have been accidental, such as the establishment of wild monk parakeets in several North American cities after their escape from captivity. Some species, including cattle egret, yellow-headed caracara and galah, have spread naturally far beyond their original ranges as agricultural expansion created alternative habitats although modern practices of intensive agriculture have negatively impacted farmland bird populations.
Anatomy and physiology
Compared with other vertebrates, birds have a body plan that shows many unusual adaptations, mostly to facilitate flight.
Skeletal system
The skeleton consists of very lightweight bones. They have large air-filled cavities (called pneumatic cavities) which connect with the respiratory system. The skull bones in adults are fused and do not show cranial sutures. The orbital cavities that house the eyeballs are large and separated from each other by a bony septum (partition). The spine has cervical, thoracic, lumbar and caudal regions with the number of cervical (neck) vertebrae highly variable and especially flexible, but movement is reduced in the anterior thoracic vertebrae and absent in the later vertebrae. The last few are fused with the pelvis to form the synsacrum. The ribs are flattened and the sternum is keeled for the attachment of flight muscles except in the flightless bird orders. The forelimbs are modified into wings. The wings are more or less developed depending on the species; the only known groups that lost their wings are the extinct moa and elephant birds. | Bird | Wikipedia | 354 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Excretory system
Like the reptiles, birds are primarily uricotelic, that is, their kidneys extract nitrogenous waste from their bloodstream and excrete it as uric acid, instead of urea or ammonia, through the ureters into the intestine. Birds do not have a urinary bladder or external urethral opening and (with exception of the ostrich) uric acid is excreted along with faeces as a semisolid waste. However, birds such as hummingbirds can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. They also excrete creatine, rather than creatinine like mammals. This material, as well as the output of the intestines, emerges from the bird's cloaca. The cloaca is a multi-purpose opening: waste is expelled through it, most birds mate by joining cloaca, and females lay eggs from it. In addition, many species of birds regurgitate pellets.
It is a common but not universal feature of altricial passerine nestlings (born helpless, under constant parental care) that instead of excreting directly into the nest, they produce a fecal sac. This is a mucus-covered pouch that allows parents to either dispose of the waste outside the nest or to recycle the waste through their own digestive system. | Bird | Wikipedia | 298 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Reproductive system
Most male birds do not have intromittent penises. Males within Palaeognathae (with the exception of the kiwis), the Anseriformes (with the exception of screamers), and in rudimentary forms in Galliformes (but fully developed in Cracidae) possess a penis, which is never present in Neoaves. Its length is thought to be related to sperm competition and it fills with lymphatic fluid instead of blood when erect. When not copulating, it is hidden within the proctodeum compartment within the cloaca, just inside the vent. Female birds have sperm storage tubules that allow sperm to remain viable long after copulation, a hundred days in some species. Sperm from multiple males may compete through this mechanism. Most female birds have a single ovary and a single oviduct, both on the left side, but there are exceptions: species in at least 16 different orders of birds have two ovaries. Even these species, however, tend to have a single oviduct. It has been speculated that this might be an adaptation to flight, but males have two testes, and it is also observed that the gonads in both sexes decrease dramatically in size outside the breeding season. Also terrestrial birds generally have a single ovary, as does the platypus, an egg-laying mammal. A more likely explanation is that the egg develops a shell while passing through the oviduct over a period of about a day, so that if two eggs were to develop at the same time, there would be a risk to survival. While rare, mostly abortive, parthenogenesis is not unknown in birds and eggs can be diploid, automictic and results in male offspring.
Birds are solely gonochoric, meaning they have two sexes: either female or male. The sex of birds is determined by the Z and W sex chromosomes, rather than by the X and Y chromosomes present in mammals. Male birds have two Z chromosomes (ZZ), and female birds have a W chromosome and a Z chromosome (WZ). A complex system of disassortative mating with two morphs is involved in the white-throated sparrow Zonotrichia albicollis, where white- and tan-browed morphs of opposite sex pair, making it appear as if four sexes were involved since any individual is compatible with only a fourth of the population. | Bird | Wikipedia | 509 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
In nearly all species of birds, an individual's sex is determined at fertilisation. However, one 2007 study claimed to demonstrate temperature-dependent sex determination among the Australian brushturkey, for which higher temperatures during incubation resulted in a higher female-to-male sex ratio. This, however, was later proven to not be the case. These birds do not exhibit temperature-dependent sex determination, but temperature-dependent sex mortality.
Respiratory and circulatory systems
Birds have one of the most complex respiratory systems of all animal groups. Upon inhalation, 75% of the fresh air bypasses the lungs and flows directly into a posterior air sac which extends from the lungs and connects with air spaces in the bones and fills them with air. The other 25% of the air goes directly into the lungs. When the bird exhales, the used air flows out of the lungs and the stored fresh air from the posterior air sac is simultaneously forced into the lungs. Thus, a bird's lungs receive a constant supply of fresh air during both inhalation and exhalation. Sound production is achieved using the syrinx, a muscular chamber incorporating multiple tympanic membranes which diverges from the lower end of the trachea; the trachea being elongated in some species, increasing the volume of vocalisations and the perception of the bird's size.
In birds, the main arteries taking blood away from the heart originate from the right aortic arch (or pharyngeal arch), unlike in the mammals where the left aortic arch forms this part of the aorta. The postcava receives blood from the limbs via the renal portal system. Unlike in mammals, the circulating red blood cells in birds retain their nucleus.
Heart type and features
The avian circulatory system is driven by a four-chambered, myogenic heart contained in a fibrous pericardial sac. This pericardial sac is filled with a serous fluid for lubrication. The heart itself is divided into a right and left half, each with an atrium and ventricle. The atrium and ventricles of each side are separated by atrioventricular valves which prevent back flow from one chamber to the next during contraction. Being myogenic, the heart's pace is maintained by pacemaker cells found in the sinoatrial node, located on the right atrium. | Bird | Wikipedia | 494 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The sinoatrial node uses calcium to cause a depolarising signal transduction pathway from the atrium through right and left atrioventricular bundle which communicates contraction to the ventricles. The avian heart also consists of muscular arches that are made up of thick bundles of muscular layers. Much like a mammalian heart, the avian heart is composed of endocardial, myocardial and epicardial layers. The atrium walls tend to be thinner than the ventricle walls, due to the intense ventricular contraction used to pump oxygenated blood throughout the body. Avian hearts are generally larger than mammalian hearts when compared to body mass. This adaptation allows more blood to be pumped to meet the high metabolic need associated with flight.
Organisation
Birds have a very efficient system for diffusing oxygen into the blood; birds have a ten times greater surface area to gas exchange volume than mammals. As a result, birds have more blood in their capillaries per unit of volume of lung than a mammal. The arteries are composed of thick elastic muscles to withstand the pressure of the ventricular contractions, and become more rigid as they move away from the heart. Blood moves through the arteries, which undergo vasoconstriction, and into arterioles which act as a transportation system to distribute primarily oxygen as well as nutrients to all tissues of the body. As the arterioles move away from the heart and into individual organs and tissues they are further divided to increase surface area and slow blood flow. Blood travels through the arterioles and moves into the capillaries where gas exchange can occur.
Capillaries are organised into capillary beds in tissues; it is here that blood exchanges oxygen for carbon dioxide waste. In the capillary beds, blood flow is slowed to allow maximum diffusion of oxygen into the tissues. Once the blood has become deoxygenated, it travels through venules then veins and back to the heart. Veins, unlike arteries, are thin and rigid as they do not need to withstand extreme pressure. As blood travels through the venules to the veins a funneling occurs called vasodilation bringing blood back to the heart. Once the blood reaches the heart, it moves first into the right atrium, then the right ventricle to be pumped through the lungs for further gas exchange of carbon dioxide waste for oxygen. Oxygenated blood then flows from the lungs through the left atrium to the left ventricle where it is pumped out to the body.
Nervous system | Bird | Wikipedia | 512 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The nervous system is large relative to the bird's size. The most developed part of the brain of birds is the one that controls the flight-related functions, while the cerebellum coordinates movement and the cerebrum controls behaviour patterns, navigation, mating and nest building. Most birds have a poor sense of smell with notable exceptions including kiwis, New World vultures and tubenoses. The avian visual system is usually highly developed. Water birds have special flexible lenses, allowing accommodation for vision in air and water. Some species also have dual fovea. Birds are tetrachromatic, possessing ultraviolet (UV) sensitive cone cells in the eye as well as green, red and blue ones. They also have double cones, likely to mediate achromatic vision.
Many birds show plumage patterns in ultraviolet that are invisible to the human eye; some birds whose sexes appear similar to the naked eye are distinguished by the presence of ultraviolet reflective patches on their feathers. Male blue tits have an ultraviolet reflective crown patch which is displayed in courtship by posturing and raising of their nape feathers. Ultraviolet light is also used in foraging—kestrels have been shown to search for prey by detecting the UV reflective urine trail marks left on the ground by rodents. With the exception of pigeons and a few other species, the eyelids of birds are not used in blinking. Instead the eye is lubricated by the nictitating membrane, a third eyelid that moves horizontally. The nictitating membrane also covers the eye and acts as a contact lens in many aquatic birds. The bird retina has a fan shaped blood supply system called the pecten.
Eyes of most birds are large, not very round and capable of only limited movement in the orbits, typically 10–20°. Birds with eyes on the sides of their heads have a wide visual field, while birds with eyes on the front of their heads, such as owls, have binocular vision and can estimate the depth of field. The avian ear lacks external pinnae but is covered by feathers, although in some birds, such as the Asio, Bubo and Otus owls, these feathers form tufts which resemble ears. The inner ear has a cochlea, but it is not a spiral as in mammals. | Bird | Wikipedia | 467 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Defence and intraspecific combat
A few species are able to use chemical defences against predators; some Procellariiformes can eject an unpleasant stomach oil against an aggressor, and some species of pitohuis from New Guinea have a powerful neurotoxin in their skin and feathers.
A lack of field observations limit our knowledge, but intraspecific conflicts are known to sometimes result in injury or death. The screamers (Anhimidae), some jacanas (Jacana, Hydrophasianus), the spur-winged goose (Plectropterus), the torrent duck (Merganetta) and nine species of lapwing (Vanellus) use a sharp spur on the wing as a weapon. The steamer ducks (Tachyeres), geese and swans (Anserinae), the solitaire (Pezophaps), sheathbills (Chionis), some guans (Crax) and stone curlews (Burhinus) use a bony knob on the alular metacarpal to punch and hammer opponents. The jacanas Actophilornis and Irediparra have an expanded, blade-like radius. The extinct Xenicibis was unique in having an elongate forelimb and massive hand which likely functioned in combat or defence as a jointed club or flail. Swans, for instance, may strike with the bony spurs and bite when defending eggs or young.
Feathers, plumage, and scales
Feathers are a feature characteristic of birds (though also present in some dinosaurs not currently considered to be true birds). They facilitate flight, provide insulation that aids in thermoregulation, and are used in display, camouflage, and signalling. There are several types of feathers, each serving its own set of purposes. Feathers are epidermal growths attached to the skin and arise only in specific tracts of skin called pterylae. The distribution pattern of these feather tracts (pterylosis) is used in taxonomy and systematics. The arrangement and appearance of feathers on the body, called plumage, may vary within species by age, social status, and sex. | Bird | Wikipedia | 445 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Plumage is regularly moulted; the standard plumage of a bird that has moulted after breeding is known as the "" plumage, or—in the Humphrey–Parkes terminology—"basic" plumage; breeding plumages or variations of the basic plumage are known under the Humphrey–Parkes system as "" plumages. Moulting is annual in most species, although some may have two moults a year, and large birds of prey may moult only once every few years. Moulting patterns vary across species. In passerines, flight feathers are replaced one at a time with the innermost being the first. When the fifth of sixth primary is replaced, the outermost begin to drop. After the innermost tertiaries are moulted, the starting from the innermost begin to drop and this proceeds to the outer feathers (centrifugal moult). The greater primary are moulted in synchrony with the primary that they overlap.
A small number of species, such as ducks and geese, lose all of their flight feathers at once, temporarily becoming flightless. As a general rule, the tail feathers are moulted and replaced starting with the innermost pair. Centripetal moults of tail feathers are however seen in the Phasianidae. The centrifugal moult is modified in the tail feathers of woodpeckers and treecreepers, in that it begins with the second innermost pair of feathers and finishes with the central pair of feathers so that the bird maintains a functional climbing tail. The general pattern seen in passerines is that the primaries are replaced outward, secondaries inward, and the tail from centre outward. Before nesting, the females of most bird species gain a bare brood patch by losing feathers close to the belly. The skin there is well supplied with blood vessels and helps the bird in incubation.
Feathers require maintenance and birds preen or groom them daily, spending an average of around 9% of their daily time on this. The bill is used to brush away foreign particles and to apply waxy secretions from the uropygial gland; these secretions protect the feathers' flexibility and act as an antimicrobial agent, inhibiting the growth of feather-degrading bacteria. This may be supplemented with the secretions of formic acid from ants, which birds receive through a behaviour known as anting, to remove feather parasites. | Bird | Wikipedia | 500 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The scales of birds are composed of the same keratin as beaks, claws, and spurs. They are found mainly on the toes and metatarsus, but may be found further up on the ankle in some birds. Most bird scales do not overlap significantly, except in the cases of kingfishers and woodpeckers.
The scales of birds are thought to be homologous to those of reptiles and mammals.
Flight
Most birds can fly, which distinguishes them from almost all other vertebrate classes. Flight is the primary means of locomotion for most bird species and is used for searching for food and for escaping from predators. Birds have various adaptations for flight, including a lightweight skeleton, two large flight muscles, the pectoralis (which accounts for 15% of the total mass of the bird) and the supracoracoideus, as well as a modified forelimb (wing) that serves as an aerofoil.
Wing shape and size generally determine a bird's flight style and performance; many birds combine powered, flapping flight with less energy-intensive soaring flight. About 60 extant bird species are flightless, as were many extinct birds. Flightlessness often arises in birds on isolated islands, most likely due to limited resources and the absence of mammalian land predators. Flightlessness is almost exclusively correlated with gigantism due to an island's inherent condition of isolation. Although flightless, penguins use similar musculature and movements to "fly" through the water, as do some flight-capable birds such as auks, shearwaters and dippers.
Behaviour
Most birds are diurnal, but some birds, such as many species of owls and nightjars, are nocturnal or crepuscular (active during twilight hours), and many coastal waders feed when the tides are appropriate, by day or night.
Diet and feeding
are varied and often include nectar, fruit, plants, seeds, carrion, and various small animals, including other birds. The digestive system of birds is unique, with a crop for storage and a gizzard that contains swallowed stones for grinding food to compensate for the lack of teeth. Some species such as pigeons and some psittacine species do not have a gallbladder. Most birds are highly adapted for rapid digestion to aid with flight. Some migratory birds have adapted to use protein stored in many parts of their bodies, including protein from the intestines, as additional energy during migration. | Bird | Wikipedia | 502 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Birds that employ many strategies to obtain food or feed on a variety of food items are called generalists, while others that concentrate time and effort on specific food items or have a single strategy to obtain food are considered specialists. Avian foraging strategies can vary widely by species. Many birds glean for insects, invertebrates, fruit, or seeds. Some hunt insects by suddenly attacking from a branch. Those species that seek pest insects are considered beneficial 'biological control agents' and their presence encouraged in biological pest control programmes. Combined, insectivorous birds eat 400–500 million metric tons of arthropods annually.
Nectar feeders such as hummingbirds, sunbirds, lories, and lorikeets amongst others have specially adapted brushy tongues and in many cases bills designed to fit co-adapted flowers. Kiwis and shorebirds with long bills probe for invertebrates; shorebirds' varied bill lengths and feeding methods result in the separation of ecological niches. Divers, diving ducks, penguins and auks pursue their prey underwater, using their wings or feet for propulsion, while aerial predators such as sulids, kingfishers and terns plunge dive after their prey. Flamingos, three species of prion, and some ducks are filter feeders. Geese and dabbling ducks are primarily grazers.
Some species, including frigatebirds, gulls, and skuas, engage in kleptoparasitism, stealing food items from other birds. Kleptoparasitism is thought to be a supplement to food obtained by hunting, rather than a significant part of any species' diet; a study of great frigatebirds stealing from masked boobies estimated that the frigatebirds stole at most 40% of their food and on average stole only 5%. Other birds are scavengers; some of these, like vultures, are specialised carrion eaters, while others, like gulls, corvids, or other birds of prey, are opportunists.
Water and drinking
Water is needed by many birds although their mode of excretion and lack of sweat glands reduces the physiological demands. Some desert birds can obtain their water needs entirely from moisture in their food. Some have other adaptations such as allowing their body temperature to rise, saving on moisture loss from evaporative cooling or panting. Seabirds can drink seawater and have salt glands inside the head that eliminate excess salt out of the nostrils. | Bird | Wikipedia | 490 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Most birds scoop water in their beaks and raise their head to let water run down the throat. Some species, especially of arid zones, belonging to the pigeon, finch, mousebird, button-quail and bustard families are capable of sucking up water without the need to tilt back their heads. Some desert birds depend on water sources and sandgrouse are particularly well known for congregating daily at waterholes. Nesting sandgrouse and many plovers carry water to their young by wetting their belly feathers. Some birds carry water for chicks at the nest in their crop or regurgitate it along with food. The pigeon family, flamingos and penguins have adaptations to produce a nutritive fluid called crop milk that they provide to their chicks.
Feather care
Feathers, being critical to the survival of a bird, require maintenance. Apart from physical wear and tear, feathers face the onslaught of fungi, ectoparasitic feather mites and bird lice. The physical condition of feathers are maintained by often with the application of secretions from the . Birds also bathe in water or dust themselves. While some birds dip into shallow water, more aerial species may make aerial dips into water and arboreal species often make use of dew or rain that collect on leaves. Birds of arid regions make use of loose soil to dust-bathe. A behaviour termed as anting in which the bird encourages ants to run through their plumage is also thought to help them reduce the ectoparasite load in feathers. Many species will spread out their wings and expose them to direct sunlight and this too is thought to help in reducing fungal and ectoparasitic activity that may lead to feather damage.
Migration
Many bird species migrate to take advantage of global differences of seasonal temperatures, therefore optimising availability of food sources and breeding habitat. These migrations vary among the different groups. Many landbirds, shorebirds, and waterbirds undertake annual long-distance migrations, usually triggered by the length of daylight as well as weather conditions. These birds are characterised by a breeding season spent in the temperate or polar regions and a non-breeding season in the tropical regions or opposite hemisphere. Before migration, birds substantially increase body fats and reserves and reduce the size of some of their organs. | Bird | Wikipedia | 461 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Migration is highly demanding energetically, particularly as birds need to cross deserts and oceans without refuelling. Landbirds have a flight range of around and shorebirds can fly up to , although the bar-tailed godwit is capable of non-stop flights of up to . Some seabirds undertake long migrations, with the longest annual migrations including those of Arctic terns, which were recorded travelling an average of between their Arctic breeding grounds in Greenland and Iceland and their wintering grounds in Antarctica, with one bird covering , and sooty shearwaters, which nest in New Zealand and Chile and make annual round trips of to their summer feeding grounds in the North Pacific off Japan, Alaska and California.
Other seabirds disperse after breeding, travelling widely but having no set migration route. Albatrosses nesting in the Southern Ocean often undertake circumpolar trips between breeding seasons.
Some bird species undertake shorter migrations, travelling only as far as is required to avoid bad weather or obtain food. Irruptive species such as the boreal finches are one such group and can commonly be found at a location in one year and absent the next. This type of migration is normally associated with food availability. Species may also travel shorter distances over part of their range, with individuals from higher latitudes travelling into the existing range of conspecifics; others undertake partial migrations, where only a fraction of the population, usually females and subdominant males, migrates. Partial migration can form a large percentage of the migration behaviour of birds in some regions; in Australia, surveys found that 44% of non-passerine birds and 32% of passerines were partially migratory.
Altitudinal migration is a form of short-distance migration in which birds spend the breeding season at higher altitudes and move to lower ones during suboptimal conditions. It is most often triggered by temperature changes and usually occurs when the normal territories also become inhospitable due to lack of food. Some species may also be nomadic, holding no fixed territory and moving according to weather and food availability. Parrots as a family are overwhelmingly neither migratory nor sedentary but considered to either be dispersive, irruptive, nomadic or undertake small and irregular migrations. | Bird | Wikipedia | 458 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The ability of birds to return to precise locations across vast distances has been known for some time; in an experiment conducted in the 1950s, a Manx shearwater released in Boston in the United States returned to its colony in Skomer, in Wales within 13 days, a distance of . Birds navigate during migration using a variety of methods. For diurnal migrants, the sun is used to navigate by day, and a stellar compass is used at night. Birds that use the sun compensate for the changing position of the sun during the day by the use of an internal clock. Orientation with the stellar compass depends on the position of the constellations surrounding Polaris. These are backed up in some species by their ability to sense the Earth's geomagnetism through specialised photoreceptors.
Communication
Birds communicate primarily using visual and auditory signals. Signals can be interspecific (between species) and intraspecific (within species).
Birds sometimes use plumage to assess and assert social dominance, to display breeding condition in sexually selected species, or to make threatening displays, as in the sunbittern's mimicry of a large predator to ward off hawks and protect young chicks.
Visual communication among birds may also involve ritualised displays, which have developed from non-signalling actions such as preening, the adjustments of feather position, pecking, or other behaviour. These displays may signal aggression or submission or may contribute to the formation of pair-bonds. The most elaborate displays occur during courtship, where "dances" are often formed from complex combinations of many possible component movements; males' breeding success may depend on the quality of such displays. | Bird | Wikipedia | 333 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Bird calls and songs, which are produced in the syrinx, are the major means by which birds communicate with sound. This communication can be very complex; some species can operate the two sides of the syrinx independently, allowing the simultaneous production of two different songs.
Calls are used for a variety of purposes, including mate attraction, evaluation of potential mates, bond formation, the claiming and maintenance of territories, the identification of other individuals (such as when parents look for chicks in colonies or when mates reunite at the start of breeding season), and the warning of other birds of potential predators, sometimes with specific information about the nature of the threat. Some birds also use mechanical sounds for auditory communication. The Coenocorypha snipes of New Zealand drive air through their feathers, woodpeckers drum for long-distance communication, and palm cockatoos use tools to drum.
Flocking and other associations
While some birds are essentially territorial or live in small family groups, other birds may form large flocks. The principal benefits of flocking are safety in numbers and increased foraging efficiency. Defence against predators is particularly important in closed habitats like forests, where ambush predation is common and multiple eyes can provide a valuable early warning system. This has led to the development of many mixed-species feeding flocks, which are usually composed of small numbers of many species; these flocks provide safety in numbers but increase potential competition for resources. Costs of flocking include bullying of socially subordinate birds by more dominant birds and the reduction of feeding efficiency in certain cases. Some species have a mixed system with breeding pairs maintaining territories, while unmated or young birds live in flocks where they secure mates prior to finding territories.
Birds sometimes also form associations with non-avian species. Plunge-diving seabirds associate with dolphins and tuna, which push shoaling fish towards the surface. Some species of hornbills have a mutualistic relationship with dwarf mongooses, in which they forage together and warn each other of nearby birds of prey and other predators.
Resting and roosting | Bird | Wikipedia | 419 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
The high metabolic rates of birds during the active part of the day is supplemented by rest at other times. Sleeping birds often use a type of sleep known as vigilant sleep, where periods of rest are interspersed with quick eye-opening "peeks", allowing them to be sensitive to disturbances and enable rapid escape from threats. Swifts are believed to be able to sleep in flight and radar observations suggest that they orient themselves to face the wind in their roosting flight. It has been suggested that there may be certain kinds of sleep which are possible even when in flight.
Some birds have also demonstrated the capacity to fall into slow-wave sleep one hemisphere of the brain at a time. The birds tend to exercise this ability depending upon its position relative to the outside of the flock. This may allow the eye opposite the sleeping hemisphere to remain vigilant for predators by viewing the outer margins of the flock. This adaptation is also known from marine mammals. Communal roosting is common because it lowers the loss of body heat and decreases the risks associated with predators. Roosting sites are often chosen with regard to thermoregulation and safety. Unusual mobile roost sites include large herbivores on the African savanna that are used by oxpeckers.
Many sleeping birds bend their heads over their backs and tuck their bills in their back feathers, although others place their beaks among their breast feathers. Many birds rest on one leg, while some may pull up their legs into their feathers, especially in cold weather. Perching birds have a tendon-locking mechanism that helps them hold on to the perch when they are asleep. Many ground birds, such as quails and pheasants, roost in trees. A few parrots of the genus Loriculus roost hanging upside down. Some hummingbirds go into a nightly state of torpor accompanied with a reduction of their metabolic rates. This physiological adaptation shows in nearly a hundred other species, including owlet-nightjars, nightjars, and woodswallows. One species, the common poorwill, even enters a state of hibernation. Birds do not have sweat glands, but can lose water directly through the skin, and they may cool themselves by moving to shade, standing in water, panting, increasing their surface area, fluttering their throat or using special behaviours like urohidrosis to cool themselves.
Breeding
Social systems | Bird | Wikipedia | 488 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
95 per cent of bird species are socially monogamous. These species pair for at least the length of the breeding season or—in some cases—for several years or until the death of one mate. Monogamy allows for both paternal care and biparental care, which is especially important for species in which care from both the female and the male parent is required in order to successfully rear a brood. Among many socially monogamous species, extra-pair copulation (infidelity) is common. Such behaviour typically occurs between dominant males and females paired with subordinate males, but may also be the result of forced copulation in ducks and other anatids.
For females, possible benefits of extra-pair copulation include getting better genes for her offspring and insuring against the possibility of infertility in her mate. Males of species that engage in extra-pair copulations will closely guard their mates to ensure the parentage of the offspring that they raise.
Other mating systems, including polygyny, polyandry, polygamy, polygynandry, and promiscuity, also occur. Polygamous breeding systems arise when females are able to raise broods without the help of males. Mating systems vary across bird families but variations within species are thought to be driven by environmental conditions. A unique system is the formation of trios where a third individual is allowed by a breeding pair temporarily into the territory to assist with brood raising thereby leading to higher fitness.
Breeding usually involves some form of courtship display, typically performed by the male. Most displays are rather simple and involve some type of song. Some displays, however, are quite elaborate. Depending on the species, these may include wing or tail drumming, dancing, aerial flights, or communal lekking. Females are generally the ones that drive partner selection, although in the polyandrous phalaropes, this is reversed: plainer males choose brightly coloured females. Courtship feeding, billing and are commonly performed between partners, generally after the birds have paired and mated.
Homosexual behaviour has been observed in males or females in numerous species of birds, including copulation, pair-bonding, and joint parenting of chicks. Over 130 avian species around the world engage in sexual interactions between the same sex or homosexual behaviours. "Same-sex courtship activities may involve elaborate displays, synchronised dances, gift-giving ceremonies, or behaviours at specific display areas including bowers, arenas, or leks."
Territories, nesting and incubation | Bird | Wikipedia | 510 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Many birds actively defend a territory from others of the same species during the breeding season; maintenance of territories protects the food source for their chicks. Species that are unable to defend feeding territories, such as seabirds and swifts, often breed in colonies instead; this is thought to offer protection from predators. Colonial breeders defend small nesting sites, and competition between and within species for nesting sites can be intense.
All birds lay amniotic eggs with hard shells made mostly of calcium carbonate. Hole and burrow nesting species tend to lay white or pale eggs, while open nesters lay camouflaged eggs. There are many exceptions to this pattern, however; the ground-nesting nightjars have pale eggs, and camouflage is instead provided by their plumage. Species that are victims of brood parasites have varying egg colours to improve the chances of spotting a parasite's egg, which forces female parasites to match their eggs to those of their hosts.
Bird eggs are usually laid in a nest. Most species create somewhat elaborate nests, which can be cups, domes, plates, mounds, or burrows. Some bird nests can be a simple scrape, with minimal or no lining; most seabird and wader nests are no more than a scrape on the ground. Most birds build nests in sheltered, hidden areas to avoid predation, but large or colonial birds—which are more capable of defence—may build more open nests. During nest construction, some species seek out plant matter from plants with parasite-reducing toxins to improve chick survival, and feathers are often used for nest insulation. Some bird species have no nests; the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body and feet. The absence of nests is especially prevalent in open habitat ground-nesting species where any addition of nest material would make the nest more conspicuous. Many ground nesting birds lay a clutch of eggs that hatch synchronously, with precocial chicks led away from the nests (nidifugous) by their parents soon after hatching. | Bird | Wikipedia | 415 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Incubation, which regulates temperature for chick development, usually begins after the last egg has been laid. In monogamous species incubation duties are often shared, whereas in polygamous species one parent is wholly responsible for incubation. Warmth from parents passes to the eggs through brood patches, areas of bare skin on the abdomen or breast of the incubating birds. Incubation can be an energetically demanding process; adult albatrosses, for instance, lose as much as of body weight per day of incubation. The warmth for the incubation of the eggs of megapodes comes from the sun, decaying vegetation or volcanic sources. Incubation periods range from 10 days (in woodpeckers, cuckoos and passerine birds) to over 80 days (in albatrosses and kiwis).
The diversity of characteristics of birds is great, sometimes even in closely related species. Several avian characteristics are compared in the table below.
Parental care and fledging
At the time of their hatching, chicks range in development from helpless to independent, depending on their species. Helpless chicks are termed altricial, and tend to be born small, blind, immobile and naked; chicks that are mobile and feathered upon hatching are termed precocial. Altricial chicks need help thermoregulating and must be brooded for longer than precocial chicks. The young of many bird species do not precisely fit into either the precocial or altricial category, having some aspects of each and thus fall somewhere on an "altricial-precocial spectrum". Chicks at neither extreme but favouring one or the other may be termed or .
The length and nature of parental care varies widely amongst different orders and species. At one extreme, parental care in megapodes ends at hatching; the newly hatched chick digs itself out of the nest mound without parental assistance and can fend for itself immediately. At the other extreme, many seabirds have extended periods of parental care, the longest being that of the great frigatebird, whose chicks take up to six months to fledge and are fed by the parents for up to an additional 14 months. The chick guard stage describes the period of breeding during which one of the adult birds is permanently present at the nest after chicks have hatched. The main purpose of the guard stage is to aid offspring to thermoregulate and protect them from predation. | Bird | Wikipedia | 506 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
In some species, both parents care for nestlings and fledglings; in others, such care is the responsibility of only one sex. In some species, other members of the same species—usually close relatives of the breeding pair, such as offspring from previous broods—will help with the raising of the young. Such alloparenting is particularly common among the Corvida, which includes such birds as the true crows, Australian magpie and fairy-wrens, but has been observed in species as different as the rifleman and red kite. Among most groups of animals, male parental care is rare. In birds, however, it is quite common—more so than in any other vertebrate class. Although territory and nest site defence, incubation, and chick feeding are often shared tasks, there is sometimes a division of labour in which one mate undertakes all or most of a particular duty.
The point at which chicks fledge varies dramatically. The chicks of the Synthliboramphus murrelets, like the ancient murrelet, leave the nest the night after they hatch, following their parents out to sea, where they are raised away from terrestrial predators. Some other species, such as ducks, move their chicks away from the nest at an early age. In most species, chicks leave the nest just before, or soon after, they are able to fly. The amount of parental care after fledging varies; albatross chicks leave the nest on their own and receive no further help, while other species continue some supplementary feeding after fledging. Chicks may also follow their parents during their first migration.
Brood parasites | Bird | Wikipedia | 331 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Brood parasitism, in which an egg-layer leaves her eggs with another individual's brood, is more common among birds than any other type of organism. After a parasitic bird lays her eggs in another bird's nest, they are often accepted and raised by the host at the expense of the host's own brood. Brood parasites may be either obligate brood parasites, which must lay their eggs in the nests of other species because they are incapable of raising their own young, or non-obligate brood parasites, which sometimes lay eggs in the nests of conspecifics to increase their reproductive output even though they could have raised their own young. One hundred bird species, including honeyguides, icterids, and ducks, are obligate parasites, though the most famous are the cuckoos. Some brood parasites are adapted to hatch before their host's young, which allows them to destroy the host's eggs by pushing them out of the nest or to kill the host's chicks; this ensures that all food brought to the nest will be fed to the parasitic chicks.
Sexual selection
Birds have evolved a variety of mating behaviours, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and colour differences are energetically costly attributes that signal competitive breeding situations. Many types of avian sexual selection have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signalling adaptations ensure that sexually selected ornaments such as plumage colouration and courtship behaviour are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviours.
Inbreeding depression
Inbreeding causes early death (inbreeding depression) in the zebra finch Taeniopygia guttata. Embryo survival (that is, hatching success of fertile eggs) was significantly lower for sib-sib mating pairs than for unrelated pairs.
Darwin's finch Geospiza scandens experiences inbreeding depression (reduced survival of offspring) and the magnitude of this effect is influenced by environmental conditions such as low food availability.
Inbreeding avoidance | Bird | Wikipedia | 500 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Incestuous matings by the purple-crowned fairy wren Malurus coronatus result in severe fitness costs due to inbreeding depression (greater than 30% reduction in hatchability of eggs). Females paired with related males may undertake extra pair matings (see Promiscuity#Other animals for 90% frequency in avian species) that can reduce the negative effects of inbreeding. However, there are ecological and demographic constraints on extra pair matings. Nevertheless, 43% of broods produced by incestuously paired females contained extra pair young.
Inbreeding depression occurs in the great tit (Parus major) when the offspring produced as a result of a mating between close relatives show reduced fitness. In natural populations of Parus major, inbreeding is avoided by dispersal of individuals from their birthplace, which reduces the chance of mating with a close relative.
Southern pied babblers Turdoides bicolor appear to avoid inbreeding in two ways. The first is through dispersal, and the second is by avoiding familiar group members as mates.
Cooperative breeding in birds typically occurs when offspring, usually males, delay dispersal from their natal group in order to remain with the family to help rear younger kin. Female offspring rarely stay at home, dispersing over distances that allow them to breed independently, or to join unrelated groups. In general, inbreeding is avoided because it leads to a reduction in progeny fitness (inbreeding depression) due largely to the homozygous expression of deleterious recessive alleles. Cross-fertilisation between unrelated individuals ordinarily leads to the masking of deleterious recessive alleles in progeny.
Ecology | Bird | Wikipedia | 344 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Birds occupy a wide range of ecological positions. While some birds are generalists, others are highly specialised in their habitat or food requirements. Even within a single habitat, such as a forest, the niches occupied by different species of birds vary, with some species feeding in the forest canopy, others beneath the canopy, and still others on the forest floor. Forest birds may be insectivores, frugivores, or nectarivores. Aquatic birds generally feed by fishing, plant eating, and piracy or kleptoparasitism. Many grassland birds are granivores. Birds of prey specialise in hunting mammals or other birds, while vultures are specialised scavengers. Birds are also preyed upon by a range of mammals including a few avivorous bats. A wide range of endo- and ectoparasites depend on birds and some parasites that are transmitted from parent to young have co-evolved and show host-specificity.
Some nectar-feeding birds are important pollinators, and many frugivores play a key role in seed dispersal. Plants and pollinating birds often coevolve, and in some cases a flower's primary pollinator is the only species capable of reaching its nectar.
Birds are often important to island ecology. Birds have frequently reached islands that mammals have not; on those islands, birds may fulfil ecological roles typically played by larger animals. For example, in New Zealand nine species of moa were important browsers, as are the kererū and kōkako today. Today the plants of New Zealand retain the defensive adaptations evolved to protect them from the extinct moa.
Many birds act as ecosystem engineers through the construction of nests, which provide important microhabitats and food for hundreds of species of invertebrates. Nesting seabirds may affect the ecology of islands and surrounding seas, principally through the concentration of large quantities of guano, which may enrich the local soil and the surrounding seas.
A wide variety of avian ecology field methods, including counts, nest monitoring, and capturing and marking, are used for researching avian ecology.
Relationship with humans | Bird | Wikipedia | 431 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Since birds are highly visible and common animals, humans have had a relationship with them since the dawn of man. Sometimes, these relationships are mutualistic, like the cooperative honey-gathering among honeyguides and African peoples such as the Borana. Other times, they may be commensal, as when species such as the house sparrow have benefited from human activities. Several species have reconciled to habits of farmers who practice traditional farming. Examples include the Sarus Crane that begins nesting in India when farmers flood the fields in anticipation of rains, and the woolly-necked storks that have taken to nesting on a short tree grown for agroforestry beside fields and canals. Several bird species have become commercially significant agricultural pests, and some pose an aviation hazard. Human activities can also be detrimental, and have threatened numerous bird species with extinction (hunting, avian lead poisoning, pesticides, roadkill, wind turbine kills and predation by pet cats and dogs are common causes of death for birds).
Birds can act as vectors for spreading diseases such as psittacosis, salmonellosis, campylobacteriosis, mycobacteriosis (avian tuberculosis), avian influenza (bird flu), giardiasis, and cryptosporidiosis over long distances. Some of these are zoonotic diseases that can also be transmitted to humans.
Economic importance
Domesticated birds raised for meat and eggs, called poultry, are the largest source of animal protein eaten by humans; in 2003, tons of poultry and tons of eggs were produced worldwide. Chickens account for much of human poultry consumption, though domesticated turkeys, ducks, and geese are also relatively common. Many species of birds are also hunted for meat. Bird hunting is primarily a recreational activity except in extremely undeveloped areas. The most important birds hunted in North and South America are waterfowl; other widely hunted birds include pheasants, wild turkeys, quail, doves, partridge, grouse, snipe, and woodcock. Muttonbirding is also popular in Australia and New Zealand. Although some hunting, such as that of muttonbirds, may be sustainable, hunting has led to the extinction or endangerment of dozens of species. | Bird | Wikipedia | 460 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Other commercially valuable products from birds include feathers (especially the down of geese and ducks), which are used as insulation in clothing and bedding, and seabird faeces (guano), which is a valuable source of phosphorus and nitrogen. The War of the Pacific, sometimes called the Guano War, was fought in part over the control of guano deposits.
Birds have been domesticated by humans both as pets and for practical purposes. Colourful birds, such as parrots and mynas, are bred in captivity or kept as pets, a practice that has led to the illegal trafficking of some endangered species. Falcons and cormorants have long been used for hunting and fishing, respectively. Messenger pigeons, used since at least 1 AD, remained important as recently as World War II. Today, such activities are more common either as hobbies, for entertainment and tourism.
Amateur bird enthusiasts (called birdwatchers, twitchers or, more commonly, birders) number in the millions. Many homeowners erect bird feeders near their homes to attract various species. Bird feeding has grown into a multimillion-dollar industry; for example, an estimated 75% of households in Britain provide food for birds at some point during the winter.
In religion and mythology
Birds play prominent and diverse roles in religion and mythology.
In religion, birds may serve as either messengers or priests and leaders for a deity, such as in the Cult of Makemake, in which the Tangata manu of Easter Island served as chiefs or as attendants, as in the case of Hugin and Munin, the two common ravens who whispered news into the ears of the Norse god Odin. In several civilisations of ancient Italy, particularly Etruscan and Roman religion, priests were involved in augury, or interpreting the words of birds while the "auspex" (from which the word "auspicious" is derived) watched their activities to foretell events. | Bird | Wikipedia | 395 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
They may also serve as religious symbols, as when Jonah (, dove) embodied the fright, passivity, mourning, and beauty traditionally associated with doves. Birds have themselves been deified, as in the case of the common peacock, which is perceived as Mother Earth by the people of southern India. In the ancient world, doves were used as symbols of the Mesopotamian goddess Inanna (later known as Ishtar), the Canaanite mother goddess Asherah, and the Greek goddess Aphrodite. In ancient Greece, Athena, the goddess of wisdom and patron deity of the city of Athens, had a little owl as her symbol. In religious images preserved from the Inca and Tiwanaku empires, birds are depicted in the process of transgressing boundaries between earthly and underground spiritual realms. Indigenous peoples of the central Andes maintain legends of birds passing to and from metaphysical worlds.
In culture and folklore
Birds have featured in culture and art since prehistoric times, when they were represented in early cave painting and carvings. Some birds have been perceived as monsters, including the mythological Roc and the Māori's legendary , a giant bird capable of snatching humans. Birds were later used as symbols of power, as in the magnificent Peacock Throne of the Mughal and Persian emperors. With the advent of scientific interest in birds, many paintings of birds were commissioned for books.
Among the most famous of these bird artists was John James Audubon, whose paintings of North American birds were a great commercial success in Europe and who later lent his name to the National Audubon Society. Birds are also important figures in poetry; for example, Homer incorporated nightingales into his Odyssey, and Catullus used a sparrow as an erotic symbol in his Catullus 2. The relationship between an albatross and a sailor is the central theme of Samuel Taylor Coleridge's The Rime of the Ancient Mariner, which led to the use of the term as a metaphor for a 'burden'. Other English metaphors derive from birds; vulture funds and vulture investors, for instance, take their name from the scavenging vulture. Aircraft, particularly military aircraft, are frequently named after birds. The predatory nature of raptors make them popular choices for fighter aircraft such as the F-16 Fighting Falcon and the Harrier Jump Jet, while the names of seabirds may be chosen for aircraft primarily used by naval forces such as the HU-16 Albatross and the V-22 Osprey. | Bird | Wikipedia | 502 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Perceptions of bird species vary across cultures. Owls are associated with bad luck, witchcraft, and death in parts of Africa, but are regarded as wise across much of Europe. Hoopoes were considered sacred in Ancient Egypt and symbols of virtue in Persia, but were thought of as thieves across much of Europe and harbingers of war in Scandinavia. In heraldry, birds, especially eagles, often appear in coats of arms In vexillology, birds are a popular choice on flags. Birds feature in the flag designs of 17 countries and numerous subnational entities and territories. Birds are used by nations to symbolise a country's identity and heritage, with 91 countries officially recognising a national bird. Birds of prey are highly represented, though some nations have chosen other species of birds with parrots being popular among smaller, tropical nations.
In music
In music, birdsong has influenced composers and musicians in several ways: they can be inspired by birdsong; they can intentionally imitate bird song in a composition, as Vivaldi, Messiaen, and Beethoven did, along with many later composers; they can incorporate recordings of birds into their works, as Ottorino Respighi first did; or like Beatrice Harrison and David Rothenberg, they can duet with birds.
A 2023 archaeological excavation of a 10,000-year-old site in Israel yielded hollow wing bones of coots and ducks with perforations made on the side that are thought to have allowed them to be used as flutes or whistles possibly used by Natufian people to lure birds of prey.
Threats and conservation
Human activities have caused population decreases or extinction in many bird species. Over a hundred bird species have gone extinct in historical times, although the most dramatic human-caused avian extinctions, eradicating an estimated 750–1800 species, occurred during the human colonisation of Melanesian, Polynesian, and Micronesian islands. Many bird populations are declining worldwide, with 1,227 species listed as threatened by BirdLife International and the IUCN in 2009. There have been long-term declines in North American bird populations, with an estimated loss of 2.9 billion breeding adults, about 30% of the total, since 1970.
The most commonly cited human threat to birds is habitat loss. Other threats include overhunting, accidental mortality due to collisions with buildings or vehicles, long-line fishing bycatch, pollution (including oil spills and pesticide use), competition and predation from nonnative invasive species, and climate change. | Bird | Wikipedia | 512 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Governments and conservation groups work to protect birds, either by passing laws that preserve and restore bird habitat or by establishing captive populations for reintroductions. Such projects have produced some successes; one study estimated that conservation efforts saved 16 species of bird that would otherwise have gone extinct between 1994 and 2004, including the California condor and Norfolk parakeet.
Human activities have allowed the expansion of a few temperate area species, such as the barn swallow and European starling. In the tropics and sub-tropics, relatively more species are expanding due to human activities, particularly due to the spread of crops such as rice whose expansion in south Asia has benefitted at least 64 bird species, though may have harmed many more species. | Bird | Wikipedia | 146 | 3410 | https://en.wikipedia.org/wiki/Bird | Biology and health sciences | Biology | null |
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a lophophore, a "crown" of tentacles used for filter feeding. Most marine bryozoans live in tropical waters, but a few are found in oceanic trenches and polar waters. The bryozoans are classified as the marine bryozoans (Stenolaemata), freshwater bryozoans (Phylactolaemata), and mostly-marine bryozoans (Gymnolaemata), a few members of which prefer brackish water. 5,869living species are known. Originally all of the crown group Bryozoa were colonial, but as an adaptation to a mesopsammal (interstitial spaces in marine sand) life or to deep-sea habitats, secondarily solitary forms have since evolved. Solitary species have been described in four genera; (Aethozooides, Aethozoon, Franzenella and Monobryozoon). The latter having a statocyst-like organ with a supposed excretory function.
The terms Polyzoa and Bryozoa were introduced in 1830 and 1831, respectively. Soon after it was named, another group of animals was discovered whose filtering mechanism looked similar, so it was included in Bryozoa until 1869, when the two groups were noted to be very different internally. The new group was given the name "Entoprocta", while the original Bryozoa were called "Ectoprocta". Disagreements about terminology persisted well into the 20th century, but "Bryozoa" is now the generally accepted term. | Bryozoa | Wikipedia | 384 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Colonies take a variety of forms, including fans, bushes and sheets. Single animals, called zooids, live throughout the colony and are not fully independent. These individuals can have unique and diverse functions. All colonies have "autozooids", which are responsible for feeding, excretion, and supplying nutrients to the colony through diverse channels. Some classes have specialist zooids like hatcheries for fertilized eggs, colonial defence structures, and root-like attachment structures. Cheilostomata is the most diverse order of bryozoan, possibly because its members have the widest range of specialist zooids. They have mineralized exoskeletons and form single-layered sheets which encrust over surfaces, and some colonies can creep very slowly by using spiny defensive zooids as legs.
Each zooid consists of a "cystid", which provides the body wall and produces the exoskeleton, and a "polypide", which holds the organs. Zooids have no special excretory organs, and autozooids' polypides are scrapped when they become overloaded with waste products; usually the body wall then grows a replacement polypide. Their gut is U-shaped, with the mouth inside the crown of tentacles and the anus outside it. Zooids of all the freshwater species are simultaneous hermaphrodites. Although those of many marine species function first as males and then as females, their colonies always contain a combination of zooids that are in their male and female stages. All species emit sperm into the water. Some also release ova into the water, while others capture sperm via their tentacles to fertilize their ova internally. In some species the larvae have large yolks, go to feed, and quickly settle on a surface. Others produce larvae that have little yolk but swim and feed for a few days before settling. After settling, all larvae undergo a radical metamorphosis that destroys and rebuilds almost all the internal tissues. Freshwater species also produce statoblasts that lie dormant until conditions are favorable, which enables a colony's lineage to survive even if severe conditions kill the mother colony. | Bryozoa | Wikipedia | 446 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Predators of marine bryozoans include sea slugs (nudibranchs), fish, sea urchins, pycnogonids, crustaceans, mites and starfish. Freshwater bryozoans are preyed on by snails, insects, and fish. In Thailand, many populations of one freshwater species have been wiped out by an introduced species of snail. Membranipora membranacea, a fast-growing invasive bryozoan off the northeast and northwest coasts of the US, has reduced kelp forests so much that it has affected local fish and invertebrate populations. Bryozoans have spread diseases to fish farms and fishermen. Chemicals extracted from a marine bryozoan species have been investigated for treatment of cancer and Alzheimer's disease, but analyses have not been encouraging.
Mineralized skeletons of bryozoans first appear in rocks from the Early Ordovician period, making it the last major phylum to appear in the fossil record. This has led researchers to suspect that bryozoans arose earlier but were initially unmineralized, and may have differed significantly from fossilized and modern forms. In 2021, some research suggested Protomelission, a genus known from the Cambrian period, could be an example of an early bryozoan, but later research suggested that this taxon may instead represent a dasyclad alga. Early fossils are mainly of erect forms, but encrusting forms gradually became dominant. It is uncertain whether the phylum is monophyletic. Bryozoans' evolutionary relationships to other phyla are also unclear, partly because scientists' view of the family tree of animals is mainly influenced by better-known phyla. Both morphological and molecular phylogeny analyses disagree over bryozoans' relationships with entoprocts, about whether bryozoans should be grouped with brachiopods and phoronids in Lophophorata, and whether bryozoans should be considered protostomes or deuterostomes.
Description
Distinguishing features | Bryozoa | Wikipedia | 428 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Bryozoans, phoronids and brachiopods strain food out of the water by means of a lophophore, a "crown" of hollow tentacles. Bryozoans form colonies consisting of clones called zooids that are typically about long. Phoronids resemble bryozoan zooids but are long and, although they often grow in clumps, do not form colonies consisting of clones. Brachiopods, generally thought to be closely related to bryozoans and phoronids, are distinguished by having shells rather like those of bivalves. All three of these phyla have a coelom, an internal cavity lined by mesothelium.
Some encrusting bryozoan colonies with mineralized exoskeletons look very like small corals. However, bryozoan colonies are founded by an ancestrula, which is round rather than shaped like a normal zooid of that species. On the other hand, the founding polyp of a coral has a shape like that of its daughter polyps, and coral zooids have no coelom or lophophore.
Entoprocts, another phylum of filter-feeders, look rather like bryozoans but their lophophore-like feeding structure has solid tentacles, their anus lies inside rather than outside the base of the "crown" and they have no coelom.
Types of zooid
All bryozoans are colonial except for one genus, Monobryozoon. Individual members of a bryozoan colony are about long and are known as zooids, since they are not fully independent animals. All colonies contain feeding zooids, known as autozooids. Those of some groups also contain non-feeding heterozooids, also known as polymorphic zooids, which serve a variety of functions other than feeding; colony members are genetically identical and co-operate, rather like the organs of larger animals. What type of zooid grows where in a colony is determined by chemical signals from the colony as a whole or sometimes in response to the scent of predators or rival colonies. | Bryozoa | Wikipedia | 438 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The bodies of all types have two main parts. The cystid consists of the body wall and whatever type of exoskeleton is secreted by the epidermis. The exoskeleton may be organic (chitin, polysaccharide or protein) or made of the mineral calcium carbonate. The latter is always absent in freshwater species. The body wall consists of the epidermis, basal lamina (a mat of non-cellular material), connective tissue, muscles, and the mesothelium which lines the coelom (main body cavity) – except that in one class, the mesothelium is split into two separate layers, the inner one forming a membranous sac that floats freely and contains the coelom, and the outer one attached to the body wall and enclosing the membranous sac in a pseudocoelom. The other main part of the bryozoan body, known as the polypide and situated almost entirely within the cystid, contains the nervous system, digestive system, some specialized muscles and the feeding apparatus or other specialized organs that take the place of the feeding apparatus.
Feeding zooids
The most common type of zooid is the feeding autozooid, in which the polypide bears a "crown" of hollow tentacles called a lophophore, which captures food particles from the water. In all colonies a large percentage of zooids are autozooids, and some consist entirely of autozooids, some of which also engage in reproduction.
The basic shape of the "crown" is a full circle. Among the freshwater bryozoans (Phylactolaemata) the crown appears U-shaped, but this impression is created by a deep dent in the rim of the crown, which has no gap in the fringe of tentacles. The sides of the tentacles bear fine hairs called cilia, whose beating drives a water current from the tips of the tentacles to their bases, where it exits. Food particles that collide with the tentacles are trapped by mucus, and further cilia on the inner surfaces of the tentacles move the particles towards the mouth in the center. The method used by ectoprocts is called "upstream collecting", as food particles are captured before they pass through the field of cilia that creates the feeding current. This method is also used by phoronids, brachiopods and pterobranchs. | Bryozoa | Wikipedia | 501 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The lophophore and mouth are mounted on a flexible tube called the "invert", which can be turned inside-out and withdrawn into the polypide, rather like the finger of a rubber glove; in this position the lophophore lies inside the invert and is folded like the spokes of an umbrella. The invert is withdrawn, sometimes within 60milliseconds, by a pair of retractor muscles that are anchored at the far end of the cystid. Sensors at the tips of the tentacles may check for signs of danger before the invert and lophophore are fully extended. Extension is driven by an increase in internal fluid pressure, which species with flexible exoskeletons produce by contracting circular muscles that lie just inside the body wall, while species with a membranous sac use circular muscles to squeeze this. Some species with rigid exoskeletons have a flexible membrane that replaces part of the exoskeleton, and transverse muscles anchored on the far side of the exoskeleton increase the fluid pressure by pulling the membrane inwards. In others there is no gap in the protective skeleton, and the transverse muscles pull on a flexible sac which is connected to the water outside by a small pore; the expansion of the sac increases the pressure inside the body and pushes the invert and lophophore out. In some species the retracted invert and lophophore are protected by an operculum ("lid"), which is closed by muscles and opened by fluid pressure. In one class, a hollow lobe called the "epistome" overhangs the mouth. | Bryozoa | Wikipedia | 341 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The gut is U-shaped, running from the mouth, in the center of the lophophore, down into the animal's interior and then back to the anus, which is located on the invert, outside and usually below the lophophore. A network of strands of mesothelium called "funiculi" ("little ropes") connects the mesothelium covering the gut with that lining the body wall. The wall of each strand is made of mesothelium, and surrounds a space filled with fluid, thought to be blood. A colony's zooids are connected, enabling autozooids to share food with each other and with any non-feeding heterozooids. The method of connection varies between the different classes of bryozoans, ranging from quite large gaps in the body walls to small pores through which nutrients are passed by funiculi.
There is a nerve ring round the pharynx (throat) and a ganglion that serves as a brain to one side of this. Nerves run from the ring and ganglion to the tentacles and to the rest of the body. Bryozoans have no specialized sense organs, but cilia on the tentacles act as sensors. Members of the genus Bugula grow towards the sun, and therefore must be able to detect light. In colonies of some species, signals are transmitted between zooids through nerves that pass through pores in the body walls, and coordinate activities such as feeding and the retraction of lophophores.
The solitary individuals of Monobryozoon are autozooids with pear-shaped bodies. The wider ends have up to 15 short, muscular projections by which the animals anchor themselves to sand or gravel and pull themselves through the sediments. | Bryozoa | Wikipedia | 361 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Avicularia and vibracula
Some authorities use the term avicularia (plural of avicularium) to refer to any type of zooid in which the lophophore is replaced by an extension that serves some protective function, while others restrict the term to those that defend the colony by snapping at invaders and small predators, killing some and biting the appendages of others. In some species the snapping zooids are mounted on a peduncle (stalk), their bird-like appearance responsible for the term – Charles Darwin described these as like "the head and beak of a vulture in miniature, seated on a neck and capable of movement". Stalked avicularia are placed upside-down on their stalks. The "lower jaws" are modified versions of the opercula that protect the retracted lophophores in autozooids of some species, and are snapped shut "like a mousetrap" by similar muscles, while the beak-shaped upper jaw is the inverted body wall. In other species the avicularia are stationary box-like zooids laid the normal way up, so that the modified operculum snaps down against the body wall. In both types the modified operculum is opened by other muscles that attach to it, or by internal muscles that raise the fluid pressure by pulling on a flexible membrane. The actions of these snapping zooids are controlled by small, highly modified polypides that are located inside the "mouth" and bear tufts of short sensory cilia. These zooids appear in various positions: some take the place of autozooids, some fit into small gaps between autozooids, and small avicularia may occur on the surfaces of other zooids.
In vibracula, regarded by some as a type of avicularia, the operculum is modified to form a long bristle that has a wide range of motion. They may function as defenses against predators and invaders, or as cleaners. In some species that form mobile colonies, vibracula around the edges are used as legs for burrowing and walking. | Bryozoa | Wikipedia | 426 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Structural polymorphs
Kenozooids (from the Greek 'empty') consist only of the body wall and funicular strands crossing the interior, and no polypide. The functions of these zooids include forming the stems of branching structures, acting as spacers that enable colonies to grow quickly in a new direction, strengthening the colony's branches, and elevating the colony slightly above its substrate for competitive advantages against other organisms. Some kenozooids are hypothesized to be capable of storing nutrients for the colony. Because kenozooids' function is generally structural, they are called "structural polymorphs."
Some heterozooids found in extinct trepostome bryozoans, called mesozooids, are thought to have functioned to space the feeding autozooids an appropriate distance apart. In thin sections of trepostome fossils, mesozooids can be seen in between the tubes that held autozooids; they are smaller tubes that are divided along their length by diaphragms, making them look like rows of box-like chambers sandwiched between autozooidal tubes.
Reproductive polymorphs
Gonozooids act as brood chambers for fertilized eggs. Almost all modern cyclostome bryozoans have them, but they can be hard to locate on a colony because there are so few gonozooids in one colony. The aperture in gonozooids, which is called an ooeciopore, acts as a point for larvae to exit. Some gonozooids have very complex shapes with autozooidal tubes passing through chambers within them. All larvae released from a gonozooid are clones created by division of a single egg; this is called monozygotic polyembryony, and is a reproductive strategy also used by armadillos.
Cheilostome bryozoans also brood their embryos; one of the common methods is through ovicells, capsules attached to autozooids. The autozooids possessing ovicells are normally still able to feed, however, so these are not considered heterozooids. | Bryozoa | Wikipedia | 440 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
"Female" polymorphs are more common than "male" polymorphs, but specialized zooids that produce sperm are also known. These are called androzooids, and some are found in colonies of Odontoporella bishopi, a species that is symbiotic with hermit crabs and lives on their shells. These zooids are smaller than the others and have four short tentacles and four long tentacles, unlike the autozooids which have 15–16 tentacles. Androzooids are also found in species with mobile colonies that can crawl around. It is possible that androzooids are used to exchange sperm between colonies when two mobile colonies or bryozoan-encrusted hermit crabs happen to encounter one another.
Other polymorphs
Spinozooids are hollow, movable spines, like very slender, small tubes, present on the surface of colonies, which probably are for defense. Some species have miniature nanozooids with small single-tentacled polypides, and these may grow on other zooids or within the body walls of autozooids that have degenerated.
Colony forms and composition
Although zooids are microscopic, colonies range in size from to over . However, the majority are under across. The shapes of colonies vary widely, depend on the pattern of budding by which they grow, the variety of zooids present and the type and amount of skeletal material they secrete.
Some marine species are bush-like or fan-like, supported by "trunks" and "branches" formed by kenozooids, with feeding autozooids growing from these. Colonies of these types are generally unmineralized but may have exoskeletons made of chitin. Others look like small corals, producing heavy lime skeletons. Many species form colonies which consist of sheets of autozooids. These sheets may form leaves, tufts or, in the genus Thalamoporella, structures that resemble an open head of lettuce. | Bryozoa | Wikipedia | 408 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The most common marine form, however, is encrusting, in which a one-layer sheet of zooids spreads over a hard surface or over seaweed. Some encrusting colonies may grow to over and contain about 2,000,000 zooids. These species generally have exoskeletons reinforced with calcium carbonate, and the openings through which the lophophores protrude are on the top or outer surface. The moss-like appearance of encrusting colonies is responsible for the phylum's name (Ancient Greek words meaning 'moss' and meaning 'animal'). Large colonies of encrusting species often have "chimneys", gaps in the canopy of lophophores, through which they swiftly expel water that has been sieved, and thus avoid re-filtering water that is already exhausted. They are formed by patches of non-feeding heterozooids. New chimneys appear near the edges of expanding colonies, at points where the speed of the outflow is already high, and do not change position if the water flow changes.
Some freshwater species secrete a mass of gelatinous material, up to in diameter, to which the zooids stick. Other freshwater species have plant-like shapes with "trunks" and "branches", which may stand erect or spread over the surface. A few species can creep at about per day.
Each colony grows by asexual budding from a single zooid known as the ancestrula, which is round rather than shaped like a normal zooid. This occurs at the tips of "trunks" or "branches" in forms that have this structure. Encrusting colonies grow round their edges. In species with calcareous exoskeletons, these do not mineralize until the zooids are fully grown. Colony lifespans range from one to about 12 years, and the short-lived species pass through several generations in one season. | Bryozoa | Wikipedia | 402 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Species that produce defensive zooids do so only when threats have already appeared, and may do so within 48 hours. The theory of "induced defenses" suggests that production of defenses is expensive and that colonies which defend themselves too early or too heavily will have reduced growth rates and lifespans. This "last minute" approach to defense is feasible because the loss of zooids to a single attack is unlikely to be significant. Colonies of some encrusting species also produce special heterozooids to limit the expansion of other encrusting organisms, especially other bryozoans. In some cases this response is more belligerent if the opposition is smaller, which suggests that zooids on the edge of a colony can somehow sense the size of the opponent. Some species consistently prevail against certain others, but most turf wars are indecisive and the combatants soon turn to growing in uncontested areas. Bryozoans competing for territory do not use the sophisticated techniques employed by sponges or corals, possibly because the shortness of bryozoan lifespans makes heavy investment in turf wars unprofitable.
Bryozoans have contributed to carbonate sedimentation in marine life since the Ordovician period. Bryozoans take responsibility for many of the colony forms, which have evolved in different taxonomic groups and vary in sediment producing ability. The nine basic bryozoan colony-forms include: encrusting, dome-shaped, palmate, foliose, fenestrate, robust branching, delicate branching, articulated and free-living. Most of these sediments come from two distinct groups of colonies: domal, delicate branching, robust branching and palmate; and fenestrate. Fenestrate colonies generate rough particles both as sediment and components of stromatoporoids coral reefs. The delicate colonies however, create both coarse sediment and form the cores of deep-water, subphotic biogenic mounds. Nearly all post- bryozoan sediments are made up of growth forms, with the addition to free-living colonies which include significant numbers of various colonies. "In contrast to the Palaeozoic, post-Palaeozoic bryozoans generated sediment varying more widely with the size of their grains; they grow as they moved from mud, to sand, to gravel."
Taxonomy | Bryozoa | Wikipedia | 477 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The phylum was originally called "Polyzoa", but this name was eventually replaced by Ehrenberg's term "Bryozoa". The name "Bryozoa" was originally applied only to the animals also known as Ectoprocta (), in which the anus lies outside the "crown" of tentacles. After the discovery of the Entoprocta (), in which the anus lies within a "crown" of tentacles, the name "Bryozoa" was promoted to phylum level to include the two classes Ectoprocta and Entoprocta. However, in 1869 Hinrich Nitsche regarded the two groups as quite distinct for a variety of reasons, and coined the name "Ectoprocta" for Ehrenberg's "Bryozoa". Despite their apparently similar methods of feeding, they differed markedly anatomically; in addition to the different positions of the anus, ectoprocts have hollow tentacles and a coelom, while entoprocts have solid tentacles and no coelom. Hence the two groups are now widely regarded as separate phyla, and the name "Bryozoa" is now synonymous with "Ectoprocta". This has remained the majority view ever since, although most publications have preferred the name "Bryozoa" rather than "Ectoprocta". Nevertheless, some notable scientists have continued to regard the "Ectoprocta" and Entoprocta as close relatives and group them under "Bryozoa".
The ambiguity about the scope of the name "Bryozoa" led to proposals in the 1960s and 1970s that it should be avoided and the unambiguous term "Ectoprocta" should be used. However, the change would have made it harder to find older works in which the phylum was called "Bryozoa", and the desire to avoid ambiguity, if applied consistently to all classifications, would have necessitated renaming of several other phyla and many lower-level groups. In practice, zoological naming of split or merged groups of animals is complex and not completely consistent. Works since 2000 have used various names to resolve the ambiguity, including: "Bryozoa", "Ectoprocta", "Bryozoa (Ectoprocta)", and "Ectoprocta (Bryozoa)". Some have used more than one approach in the same work. | Bryozoa | Wikipedia | 511 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The common name "moss animals" is the literal meaning of "Bryozoa", from Greek ('moss') and ('animals'), based on the mossy appearance of encrusting species.
Until 2008 there were "inadequately known and misunderstood type species belonging to the Cyclostome Bryozoan family Oncousoeciidae." Modern research and experiments have been done using low-vacuum scanning electron microscopy of uncoated type material to critically examine and perhaps revise the taxonomy of three genera belonging to this family, including Oncousoecia, Microeciella, and Eurystrotos. This method permits data to be obtained that would be difficult to recognize with an optical microscope. The valid type species of Oncousoecia was found to be Oncousoecia lobulata. This interpretation stabilizes Oncousoecia by establishing a type species that corresponds to the general usage of the genus. Fellow Oncousoeciid Eurystrotos is now believed to be not conspecific with O. lobulata, as previously suggested, but shows enough similarities to be considered a junior synonym of Oncousoecia. Microeciella suborbicularus has also been recently distinguished from O. lobulata and O. dilatans, using this modern method of low vacuum scanning, with which it has been inaccurately synonymized with in the past. A new genus has also been recently discovered called Junerossia in the family Stomachetosellidae, along with 10 relatively new species of bryozoa such as Alderina flaventa, Corbulella extenuata, Puellina septemcryptica, Junerossia copiosa, Calyptotheca kapaaensis, Bryopesanser serratus, Cribellopora souleorum, Metacleidochasma verrucosa, Disporella compta, and Favosipora adunca.
Classification and diversity
Counts of formally described species range between 4,000 and 4,500. The Gymnolaemata and especially Cheilostomata have the greatest numbers of species, possibly because of their wide range of specialist zooids. Under the Linnaean system of classification, which is still used as a convenient way to label groups of organisms, living members of the phylum Bryozoa are divided into:
Fossil record | Bryozoa | Wikipedia | 496 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Fossils of about 15,000 bryozoan species have been found. Bryozoans are among the three dominant groups of Paleozoic fossils. Bryozoans with calcitic skeletons were a major source of the carbonate minerals that make up limestones, and their fossils are incredibly common in marine sediments worldwide from the Ordovician onward. However, unlike corals and other colonial animals found in the fossil record, Bryozoan colonies did not reach large sizes. Fossil bryozoan colonies are typically found highly fragmented and scattered; the preservation of complete zoaria is uncommon in the fossil record, and relatively little study has been devoted to reassembling fragmented zoaria. The largest known fossil colonies are branching trepostome bryozoans from Ordovician rocks in the United States, reaching 66 centimeters in height. | Bryozoa | Wikipedia | 173 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
The oldest species with a mineralized skeleton occurs in the Lower Ordovician. It is likely that the first bryozoans appeared much earlier and were entirely soft-bodied, and the Ordovician fossils record the appearance of mineralized skeletons in this phylum. By the Arenigian stage of the Early Ordovician period, about , all the modern orders of stenolaemates were present, and the ctenostome order of gymnolaemates had appeared by the Middle Ordovician, about . The Early Ordovician fossils may also represent forms that had already become significantly different from the original members of the phylum. Ctenostomes with phosphatized soft tissue are known from the Devonian. Other types of filter feeders appeared around the same time, which suggests that some change made the environment more favorable for this lifestyle. Fossils of cheilostomates, an order of gymnolaemates with mineralized skeletons, first appear in the Mid Jurassic, about , and these have been the most abundant and diverse bryozoans from the Cretaceous to the present. Evidence compiled from the last 100 million years show that cheilostomatids consistently grew over cyclostomatids in territorial struggles, which may help to explain how cheilostomatids replaced cyclostomatids as the dominant marine bryozoans. Marine fossils from the Paleozoic era, which ended , are mainly of erect forms, those from the Mesozoic are fairly equally divided by erect and encrusting forms, and more recent ones are predominantly encrusting. Fossils of the soft, freshwater phylactolaemates are very rare, appear in and after the Late Permian (which began about ) and consist entirely of their durable statoblasts. There are no known fossils of freshwater members of other classes.
Evolutionary family tree
Scientists are divided about whether the Bryozoa (Ectoprocta) are a monophyletic group (whether they include all and only a single ancestor species and all its descendants), about what are the phylum's closest relatives in the family tree of animals, and even about whether they should be regarded as members of the protostomes or deuterostomes, the two major groups that account for all moderately complex animals. | Bryozoa | Wikipedia | 483 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Molecular phylogeny, which attempts to work out the evolutionary family tree of organisms by comparing their biochemistry and especially their genes, has done much to clarify the relationships between the better-known invertebrate phyla. However, the shortage of genetic data about "minor phyla" such as bryozoans and entoprocts has left their relationships to other groups unclear.
Traditional view
The traditional view is that the Bryozoa are a monophyletic group, in which the class Phylactolaemata is most closely related to Stenolaemata and Ctenostomatida, the classes that appear earliest in the fossil record. However, in 2005 a molecular phylogeny study that focused on phylactolaemates concluded that these are more closely related to the phylum Phoronida, and especially to the only phoronid species that is colonial, than they are to the other ectoproct classes. That implies that the Entoprocta are not monophyletic, as the Phoronida are a sub-group of ectoprocts but the standard definition of Entoprocta excludes the Phoronida.
In 2009 another molecular phylogeny study, using a combination of genes from mitochondria and the cell nucleus, concluded that Bryozoa is a monophyletic phylum, in other words includes all the descendants of a common ancestor that is itself a bryozoan. The analysis also concluded that the classes Phylactolaemata, Stenolaemata and Gymnolaemata are also monophyletic, but could not determine whether Stenolaemata are more closely related to Phylactolaemata or Gymnolaemata. The Gymnolaemata are traditionally divided into the soft-bodied Ctenostomatida and mineralized Cheilostomata, but the 2009 analysis considered it more likely that neither of these orders is monophyletic and that mineralized skeletons probably evolved more than once within the early Gymnolaemata. | Bryozoa | Wikipedia | 431 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Bryozoans' relationships with other phyla are uncertain and controversial. Traditional phylogeny, based on anatomy and on the development of the adult forms from embryos, has produced no enduring consensus about the position of ectoprocts. Attempts to reconstruct the family tree of animals have largely ignored ectoprocts and other "minor phyla", which have received little scientific study because they are generally tiny, have relatively simple body plans, and have little impact on human economies – despite the fact that the "minor phyla" include most of the variety in the evolutionary history of animals.
In the opinion of Ruth Dewel, Judith Winston, and Frank McKinney, "Our standard interpretation of bryozoan morphology and embryology is a construct resulting from over 100 years of attempts to synthesize a single framework for all invertebrates," and takes little account of some peculiar features of ectoprocts.
In ectoprocts, all of the larva's internal organs are destroyed during the metamorphosis to the adult form and the adult's organs are built from the larva's epidermis and mesoderm, while in other bilaterians some organs including the gut are built from endoderm. In most bilaterian embryos the blastopore, a dent in the outer wall, deepens to become the larva's gut, but in ectoprocts the blastopore disappears and a new dent becomes the point from which the gut grows. The ectoproct coelom is formed by neither of the processes used by other bilaterians, enterocoely, in which pouches that form on the wall of the gut become separate cavities, nor schizocoely, in which the tissue between the gut and the body wall splits, forming paired cavities.
Entoprocts
When entoprocts were discovered in the 19th century, they and bryozoans (ectoprocts) were regarded as classes within the phylum Bryozoa, because both groups were sessile animals that filter-fed by means of a crown of tentacles that bore cilia. | Bryozoa | Wikipedia | 446 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
From 1869 onwards increasing awareness of differences, including the position of the entoproct anus inside the feeding structure and the difference in the early pattern of division of cells in their embryos, caused scientists to regard the two groups as separate phyla, and "Bryozoa" became just an alternative name for ectoprocts, in which the anus is outside the feeding organ. A series of molecular phylogeny studies from 1996 to 2006 have also concluded that bryozoans (ectoprocts) and entoprocts are not sister groups.
However, two well-known zoologists, Claus Nielsen and Thomas Cavalier-Smith, maintain on anatomical and developmental grounds that bryozoans and entoprocts are member of the same phylum, Bryozoa. A molecular phylogeny study in 2007 also supported this old idea, while its conclusions about other phyla agreed with those of several other analyses.
Grouping into the Lophophorata
By 1891 bryozoans (ectoprocts) were grouped with phoronids in a super-phylum called "Tentaculata". In the 1970s comparisons between phoronid larvae and the cyphonautes larva of some gymnolaete bryozoans produced suggestions that the bryozoans, most of which are colonial, evolved from a semi-colonial species of phoronid. Brachiopods were also assigned to the "Tentaculata", which were renamed Lophophorata as they all use a lophophore for filter feeding.
The majority of scientists accept this, but Claus Nielsen thinks these similarities are superficial. The Lophophorata are usually defined as animals with a lophophore, a three-part coelom and a U-shaped gut. In Nielsen's opinion, phoronids' and brachiopods' lophophores are more like those of pterobranchs, which are members of the phylum Hemichordata. Bryozoan's tentacles bear cells with multiple cilia, while the corresponding cells of phoronids', brachiopods' and pterobranchs' lophophores have one cilium per cell; and bryozoan tentacles have no hemal canal ("blood vessel"), which those of the other three phyla have. | Bryozoa | Wikipedia | 494 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
If the grouping of bryozoans with phoronids and brachiopods into Lophophorata is correct, the next issue is whether the Lophophorata are protostomes, along with most invertebrate phyla, or deuterostomes, along with chordates, hemichordates and echinoderms.
The traditional view was that lophophorates were a mix of protostome and deuterostome features. Research from the 1970s onwards suggested they were deuterostomes, because of some features that were thought characteristic of deuterostomes: a three-part coelom; radial rather than spiral cleavage in the development of the embryo; and formation of the coelom by enterocoely. However the coelom of ectoproct larvae shows no sign of division into three sections, and that of adult ectoprocts is different from that of other coelomate phyla as it is built anew from epidermis and mesoderm after metamorphosis has destroyed the larval coelom.
Lophophorate molecular phylogenetics
Molecular phylogeny analyses from 1995 onwards, using a variety of biochemical evidence and analytical techniques, placed the lophophorates as protostomes and closely related to annelids and molluscs in a super-phylum called Lophotrochozoa. "Total evidence" analyses, which used both morphological features and a relatively small set of genes, came to various conclusions, mostly favoring a close relationship between lophophorates and Lophotrochozoa. A study in 2008, using a larger set of genes, concluded that the lophophorates were closer to the Lophotrochozoa than to deuterostomes, but also that the lophophorates were not monophyletic. Instead, it concluded that brachiopods and phoronids formed a monophyletic group, but bryozoans (ectoprocts) were closest to entoprocts, supporting the original definition of "Bryozoa".
They are the only major phylum of exclusively clonal animals, composed of modular units known as zooids. Because they thrive in colonies, colonial growth allows them to develop unrestricted variations in form. Despite this, only a small number of basic growth forms have been found and have commonly reappeared throughout the history of the bryozoa. | Bryozoa | Wikipedia | 511 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
Ectoproct molecular phylogenetics
The phylogenetic position of the ectoproct bryozoans remains uncertain, but it remains certain that they belong to the Protostomia and more specifically to the Lophotrochozoa. This implies that the ectoproct larva is a trochophore with the corona being a homologue of the prototroch; this is supported from the similarity between the coronate larvae and the Type 1 pericalymma larvae of some molluscs and sipunculans, where the prototroch zone is expanded to cover the hyposphere.
A study of the mitochondrial DNA sequence suggests that the Bryozoa may be related to the Chaetognatha.
Physiology
Feeding and excretion
Most species are filter feeders that sieve small particles, mainly phytoplankton (microscopic floating plants), out of the water. The freshwater species Plumatella emarginata feeds on diatoms, green algae, cyanobacteria, non-photosynthetic bacteria, dinoflagellates, rotifers, protozoa, small nematodes, and microscopic crustaceans. While the currents that bryozoans generate to draw food towards the mouth are well understood, the exact method of capture is still debated. All species also flick larger particles towards the mouth with a tentacle, and a few capture zooplankton (planktonic animals) by using their tentacles as cages. In addition the tentacles, whose surface area is increased by microvilli (small hairs and pleats), absorb organic compounds dissolved in the water. Unwanted particles may be flicked away by tentacles or shut out by closing the mouth. A study in 2008 showed that both encrusting and erect colonies fed more quickly and grew faster in gentle than in strong currents.
In some species the first part of the stomach forms a muscular gizzard lined with chitinous teeth that crush armored prey such as diatoms. Wave-like peristaltic contractions move the food through the stomach for digestion. The final section of the stomach is lined with cilia (minute hairs) that compress undigested solids, which then pass through the intestine and out through the anus. | Bryozoa | Wikipedia | 465 | 3416 | https://en.wikipedia.org/wiki/Bryozoa | Biology and health sciences | Lophotrochozoa | null |
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