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The Communist Party was forced to give up its political monopoly on 10 November 1989 under the influence of the Revolutions of 1989. Zhivkov resigned and Bulgaria embarked on a transition to a parliamentary democracy. The first free elections in June 1990 were won by the Communist Party, now rebranded as the Bulgarian Socialist Party. A new constitution that provided for a relatively weak elected president and for a prime minister accountable to the legislature was adopted in July 1991. The new system initially failed to improve living standards or create economic growth—the average quality of life and economic performance remained lower than under communism well into the early 2000s. After 2001, economic, political and geopolitical conditions improved greatly, and Bulgaria achieved high Human Development status in 2003. It became a member of NATO in 2004 and participated in the War in Afghanistan. After several years of reforms, it joined the European Union and the single market in 2007, despite EU concerns over government corruption. Bulgaria hosted the 2018 Presidency of the Council of the European Union at the National Palace of Culture in Sofia.
Geography. Bulgaria is a middle-sized country situated in Southeast Europe, in the east of the Balkans. Its territory covers an area of , while land borders with its five neighbouring countries run a total length of , and its coastline is long. Bulgaria's geographic coordinates are 43° N 25° E. The most notable topographical features of the country are the Danubian Plain, the Balkan Mountains, the Upper Thracian Plain, and the Rila-Rhodope massif. The southern edge of the Danubian Plain slopes upward into the foothills of the Balkans, while the Danube defines the border with Romania. The Thracian Plain is roughly triangular, beginning southeast of Sofia and broadening as it reaches the Black Sea coast. The Balkan mountains run laterally through the middle of the country from west to east. The mountainous southwest has two distinct alpine type ranges—Rila and Pirin, which border the lower but more extensive Rhodope Mountains to the east, and various medium altitude mountains to west, northwest and south, like Vitosha, Osogovo and Belasitsa. Musala, at , is the highest point in both Bulgaria and the Balkans. The Black Sea coast is the country's lowest point. Plains occupy about one third of the territory, while plateaux and hills occupy 41%. Most rivers are short and with low water levels. The longest river located solely in Bulgarian territory, the Iskar, has a length of . The Struma and the Maritsa are two major rivers in the south.
Climate. Bulgaria has a varied and changeable climate, which results from being positioned at the meeting point of the Mediterranean, Oceanic and Continental air masses combined with the barrier effect of its mountains. Northern Bulgaria averages cooler, and registers more precipitation, than the regions south of the Balkan mountains. Temperature amplitudes vary significantly in different areas. The lowest recorded temperature is , while the highest is . Precipitation averages about per year, and varies from in Dobrudja to more than in the mountains. Continental air masses bring significant amounts of snowfall during winter. Considering its relatively small area, Bulgaria has variable and complex climate. The country occupies the southernmost part of the continental climatic zone, with small areas in the south falling within the Mediterranean climatic zone. The continental zone is predominant, because continental air masses flow easily into the unobstructed Danubian Plain. The continental influence, stronger during the winter, produces abundant snowfall; the Mediterranean influence increases during the second half of summer and produces hot and dry weather. Bulgaria is subdivided into five climatic zones: continental zone (Danubian Plain, Pre-Balkan and the higher valleys of the Transitional geomorphological region); transitional zone (Upper Thracian Plain, most of the Struma and Mesta valleys, the lower Sub-Balkan valleys); continental-Mediterranean zone (the southernmost areas of the Struma and Mesta valleys, the eastern Rhodope Mountains, Sakar and Strandzha); Black Sea zone along the coastline with an average length of 30–40 km inland; and alpine zone in the mountains above 1000 m altitude (central Balkan Mountains, Rila, Pirin, Vitosha, western Rhodope Mountains, etc.).
Biodiversity and conservation. The interaction of climatic, hydrological, geological and topographical conditions has produced a relatively wide variety of plant and animal species. Bulgaria's biodiversity, one of the richest in Europe, is conserved in three national parks, 11 nature parks, 10 biosphere reserves and 565 protected areas. Ninety-three of the 233 mammal species of Europe are found in Bulgaria, along with 49% of butterfly and 30% of vascular plant species. Overall, 41,493 plant and animal species are present. Larger mammals with sizable populations include deer (106,323 individuals), wild boar (88,948), golden jackal (47,293) and red fox (32,326). Partridges number some 328,000 individuals, making them the most widespread gamebird. A third of all nesting birds in Bulgaria can be found in Rila National Park, which also hosts Arctic and alpine species at high altitudes. Flora includes more than 3,800 vascular plant species of which 170 are endemic and 150 are considered endangered. A checklist of larger fungi in Bulgaria by the Institute of Botany identifies more than 1,500 species.
A checklist of larger fungi in Bulgaria by the Institute of Botany identifies more than 1,500 species. In Bulgaria forest cover is around 36% of the total land area, equivalent to 3,893,000 hectares (ha) of forest in 2020, up from 3,327,000 hectares (ha) in 1990. In 2020, naturally regenerating forest covered 3,116,000 hectares (ha) and planted forest covered 777,000 hectares (ha). Of the naturally regenerating forest 18% was reported to be primary forest (consisting of native tree species with no clearly visible indications of human activity) and around 18% of the forest area was found within protected areas. For the year 2015, 88% of the forest area was reported to be under public ownership and 12% private ownership.In 1998, the Bulgarian government adopted the National Biological Diversity Conservation Strategy, a comprehensive programme seeking the preservation of local ecosystems, protection of endangered species and conservation of genetic resources. Bulgaria has some of the largest Natura 2000 areas in Europe covering 33.8% of its territory. protection of endangered species and conservation of genetic resources. Bulgaria has some of the largest Natura 2000 areas in Europe covering 33.8% of its territory. It also achieved its Kyoto Protocol objective of reducing carbon dioxide emissions by 30% from 1990 to 2009.
Bulgaria ranks 37th in the 2024 Environmental Performance Index, but scores low on air quality. Particulate levels are the highest in Europe, especially in urban areas affected by automobile traffic and coal-based power stations. One of these, the lignite-fired Maritsa Iztok-2 station, is causing the highest damage to health and the environment in the European Union. Pesticide use in agriculture and antiquated industrial sewage systems produce extensive soil and water pollution. Water quality began to improve in 1998 and has maintained a trend of moderate improvement. Over 75% of surface rivers meet European standards for good quality. Politics. Bulgaria is a parliamentary democracy where the prime minister is the head of government and the most powerful executive position. The political system has three branches—legislative, executive and judicial, with universal suffrage for citizens at least 18 years old. The Constitution also provides possibilities of direct democracy, namely petitions and national referendums. Elections are supervised by an independent Central Election Commission that includes members from all major political parties. Parties must register with the commission prior to participating in a national election. Normally, the prime minister-elect is the leader of the party receiving the most votes in parliamentary elections, although this is not always the case.
Unlike the prime minister, presidential domestic power is more limited. The directly elected president serves as head of state and commander-in-chief of the armed forces, and has the authority to return a bill for further debate, although the parliament can override the presidential veto by a simple majority vote. Political parties gather in the National Assembly, a body of 240 deputies elected to four-year terms by direct popular vote. The National Assembly has the power to enact laws, approve the budget, schedule presidential elections, select and dismiss the prime minister and other ministers, declare war, deploy troops abroad, and ratify international treaties and agreements. Overall, Bulgaria displays a pattern of unstable governments. Boyko Borisov, the leader of the centre-right, pro-EU party GERB, served three terms as prime minister between 2009 and 2021. It won the 2009 general election and formed a minority government, which resigned in February 2013 after nationwide protests over the low living standards, corruption and the perceived failure of the democratic system.
The subsequent snap elections in May resulted in a narrow win for GERB, but the Bulgarian Socialist Party eventually formed a government led by Plamen Oresharski after Borisov failed to secure parliamentary support. The Oresharski government resigned in July 2014 amid continuing large-scale protests. The October 2014 elections resulted in a third GERB victory. Borisov formed a coalition with several right-wing parties, but resigned again after the candidate backed by his party failed to win the 2016 Presidential election. The March 2017 snap election was again won by GERB, but with 95 seats in Parliament. They formed a coalition with the far-right United Patriots, who held 27 seats. Borisov's last cabinet saw a dramatic decrease in freedom of the press, and a number of corruption revelations that triggered yet another wave of mass protests in 2020. GERB came out first in the regular April 2021 election, but with its weakest result so far. All other parties refused to form a government, and after a brief deadlock, another election was called for July 2021. It too failed to break the stalemate, as no political party was able to form a coalition government.
In April 2023, because of the political deadlock, Bulgaria held its fifth parliamentary election since April 2021. GERB was the biggest, winning 69 seats. The bloc led by We Continue the Change won 64 seats in the 240-seat parliament. In June 2023, Prime Minister Nikolai Denkov formed a new coalition between We Continue The Change and GERB. According to the coalition agreement, Denkov will lead the government for the first nine months. He will be succeeded by former European Commissioner, Mariya Gabriel, of the GERB party. She was supposed to take over as prime minister in nine months. Denkov resigned in accordance with the rotation agreement on 5 March 2024, to allow Gabriel to become the new Prime Minister. On 20 March 2024, the planned government rotation and signing of a renewed government failed due to disagreements between the two alliances on the cabinet and breaking down of relations. Negotiations ensued across to form a new government, but failed to produce any governments that could reach a majority support. GERB rejected the chance to form a government. PP–DB made a limited attempt to respect the original rotation agreement. The final chance to form a government, chosen by president Rumen Radev, went to ITN, which was immediately rejected.
On 29 March, as per Article 98 of the constitution, the President appointed the Chairman of the National Audit Office, Dimitar Glavchev, as the candidate for caretaker prime minister. He was granted a one week deadline of until 6 April 2024 to propose the composition of the caretaker government. Glavchev presented his proposal for the caretaker government on 5 April 2024, accepted by the President following negotiations and scheduling the election for 9 June 2024. Glavchev and his cabinet were inaugurated on 9 April 2024 by the National Assembly. The new elected 51st Parliament replaced the 50th Parliament when all elected members were sworn in on 11 November 2024. After 11 voting rounds, Natalia Kiselova (BSP–OL) was elected as speaker of the National Assembly on 6 December 2024. President Rumen Radev granted the first negotiation mandate to the largest party GERB-SDS on 15 January 2025, which formed a minority government alongside BSP and ITN, with support from APS (Dogan). The government is led by GERB politician Rosen Zhelyazkov.
Freedom House has reported a continuing deterioration of democratic governance after 2009, citing reduced media independence, stalled reforms, abuse of authority at the highest level and increased dependence of local administrations on the central government. Bulgaria is still listed as "Free", with a political system designated as a semi-consolidated democracy, albeit with deteriorating scores. The Democracy Index defines it as a "Flawed democracy". A 2018 survey by the Institute for Economics and Peace reported that less than 15% of respondents considered elections to be fair. Legal system. Bulgaria has a civil law legal system. The judiciary is overseen by the Ministry of Justice. The Supreme Administrative Court and the Supreme Court of Cassation are the highest courts of appeal and oversee the application of laws in subordinate courts. The Supreme Judicial Council manages the system and appoints judges. The legal system is regarded by both domestic and international observers as one of Europe's most inefficient due to a pervasive lack of transparency and corruption. Law enforcement is carried out by organisations mainly subordinate to the Ministry of the Interior. The General Directorate of National Police (GDNP) combats general crime and maintains public order. GDNP fields 26,578 police officers in its local and national sections. The bulk of criminal cases are transport-related, followed by theft and drug-related crime; homicide rates are low. The Ministry of the Interior also heads the Border Police Service and the National Gendarmerie—a specialised branch for anti-terrorist activity, crisis management and riot control. Counterintelligence and national security are the responsibility of the State Agency for National Security.
Administrative divisions. Bulgaria is a unitary state. Since the 1880s, the number of territorial management units has varied from seven to 26. Between 1987 and 1999, the administrative structure consisted of nine provinces ("oblasti", singular "oblast"). A new administrative structure was adopted in parallel with the decentralisation of the economic system. It includes 27 provinces and a metropolitan capital province (Sofia City). All areas take their names from their respective capital cities. The provinces are subdivided into 265 municipalities. Municipalities are run by mayors, who are elected to four-year terms, and by directly elected municipal councils. Bulgaria is a highly centralised state where the Council of Ministers directly appoints regional governors and all provinces and municipalities are heavily dependent on it for funding. Foreign relations. Bulgaria became a member of the United Nations in 1955. Since 1966, it has been a non-permanent member of the Security Council three times, most recently from 2002 to 2003. It was also among the founding nations of the Organisation for Security and Co-operation in Europe (OSCE) in 1975. Euro-Atlantic integration has been a priority since the fall of communism, although the communist leadership also had aspirations of leaving the Warsaw Pact and joining the European Communities by 1987. Bulgaria signed the European Union Treaty of Accession on 25 April 2005, and became a full member of the European Union on 1 January 2007. In addition, it has a tripartite economic and diplomatic collaboration with Romania and Greece, good ties with China and Vietnam and a historical relationship with Russia.
Bulgaria deployed significant numbers of both civilian and military advisors in Soviet-allied countries like Nicaragua and Libya during the Cold War. The first deployment of foreign troops on Bulgarian soil since World War II occurred in 2001, when the country hosted six KC-135 Stratotanker aircraft and 200 support personnel for the war effort in Afghanistan. International military relations were further expanded with accession to NATO in March 2004 and the US-Bulgarian Defence Cooperation Agreement signed in April 2006. Bezmer and Graf Ignatievo air bases, the Novo Selo training range, and a logistics centre in Aytos subsequently became joint military training facilities cooperatively used by the United States and Bulgarian militaries. Despite its active international defence collaborations, Bulgaria ranks as among the most peaceful countries globally, tying 6th alongside Iceland regarding domestic and international conflicts, and 26th on average in the Global Peace Index. Following the 2022 Russian invasion of Ukraine, Bulgaria decided to assist Ukraine; in 2023, after Gazprom illegally stopped exporting gas to Bulgaria, the country in turn stopped importing Russian oil and gas.
Military. The Bulgarian Armed Forces are the military of Bulgaria and are composed of land forces, navy and an air force. The Armed Forces have 36,950 active troops, supplemented by 3,000 reservists. The land forces consist of two mechanised brigades and eight independent regiments and battalions; the air force operates 106 aircraft and air defence systems across six air bases, and the navy operates various ships, helicopters and coastal defence weapons. Military inventory mainly consists of Soviet equipment like Mikoyan MiG-29 and Sukhoi Su-25 jets, S-300PT air defence systems and SS-21 Scarab short-range ballistic missiles. The Armed Forces are modernising with F-16 Block 70 fighter jets, new multi-purpose corvettes and other modern NATO-standard equipment. Bulgaria is in the process of buying new US-built Stryker vehicles, new 155 mm self-propelled howitzers, new 3D early-warning radars, new surface-to-air missiles and more. Economy. Bulgaria has an open, high-income range market economy where the private sector accounts for more than 70% of GDP. From a largely agricultural country with a predominantly rural population in 1948, by the 1980s Bulgaria had transformed into an industrial economy, with scientific and technological research at the top of its budgetary expenditure priorities. The loss of COMECON markets in 1990 and the subsequent "shock therapy" of the planned system caused a steep decline in industrial and agricultural production, ultimately followed by an economic collapse in 1997. The economy largely recovered during a period of rapid growth several years later, but the average salary of 2,072 leva ($1,142) per month remains the lowest in the EU.
A balanced budget was achieved in 2003 and the country began running a surplus the following year. Expenditures amounted to $21.15 billion and revenues were $21.67 billion in 2017. Most government spending on institutions is earmarked for security. The ministries of defence, the interior and justice are allocated the largest share of the annual government budget, whereas those responsible for the environment, tourism and energy receive the least funding. Taxes form the bulk of government revenue at 30% of GDP. Bulgaria has some of the lowest corporate income tax rates in the EU at a flat 10% rate. The tax system is two-tier. Value added tax, excise duties, corporate and personal income tax are national, whereas real estate, inheritance, and vehicle taxes are levied by local authorities. Strong economic performance in the early 2000s reduced government debt from 79.6% in 1998 to 14.1% in 2008. It has since increased to 22.6% of GDP by 2022, but remains the second lowest in the EU. The Yugozapaden planning area is the most developed region with a per capita gross domestic product (PPP) of $29,816 in 2018. It includes the capital city and the surrounding Sofia Province, which alone generate 42% of national gross domestic product despite hosting only 22% of the population. GDP per capita (in PPS) and the cost of living in 2019 stood at 53 and 52.8% of the EU average (100%), respectively. National PPP GDP was estimated at $143.1 billion in 2016, with a per capita value of $20,116. Economic growth statistics take into account illegal transactions from the informal economy, which is the largest in the EU as a percentage of economic output. The Bulgarian National Bank issues the national currency, lev, which is pegged to the euro at a rate of 1.95583 levа per euro.
After several consecutive years of high growth, repercussions of the 2008 financial crisis resulted in a 3.6% contraction of GDP in 2009 and increased unemployment. Positive growth was restored in 2010 but intercompany debt exceeded $59 billion, meaning that 60% of all Bulgarian companies were mutually indebted. By 2012, it had increased to $97 billion, or 227% of GDP. The government implemented strict austerity measures with IMF and EU encouragement to some positive fiscal results, but the social consequences of these measures, such as increased income inequality and accelerated outward migration, have been "catastrophic" according to the International Trade Union Confederation. Siphoning of public funds to the families and relatives of politicians from incumbent parties has resulted in fiscal and welfare losses to society. Bulgaria ranks 71st in the Corruption Perceptions Index and experiences the worst levels of corruption in the European Union, a phenomenon that remains a source of profound public discontent. Along with organised crime, corruption has resulted in a rejection of the country's Schengen Area application and withdrawal of foreign investment, though the country officially became a full member of the zone in January 2025. Government officials reportedly engage in embezzlement, influence trading, government procurement violations and bribery with impunity. Government procurement in particular is a critical area in corruption risk. An estimated 10 billion leva ($5.99 billion) of state budget and European cohesion funds are spent on public tenders each year; nearly 14 billion ($8.38 billion) were spent on public contracts in 2017 alone. A large share of these contracts are awarded to a few politically connected companies amid widespread irregularities, procedure violations and tailor-made award criteria. Despite repeated criticism from the European Commission, EU institutions refrain from taking measures against Bulgaria because it supports Brussels on a number of issues, unlike Poland or Hungary.
Structure and sectors. The labour force is 3.36 million people, of whom 6.8% are employed in agriculture, 26.6% in industry and 66.6% in the services sector. Extraction of metals and minerals, production of chemicals, machine building, steel, biotechnology, tobacco, food processing and petroleum refining are among the major industrial activities. Mining alone employs 24,000 people and generates about 5% of the country's GDP; the number of employed in all mining-related industries is 120,000. Bulgaria is Europe's fifth-largest coal producer. Local deposits of coal, iron, copper and lead are vital for the manufacturing and energy sectors. The main destinations of Bulgarian exports outside the EU are Turkey, China and Serbia, while Russia, Turkey and China are by far the largest import partners. Most of the exports are manufactured goods, machinery, chemicals, fuel products and food. Two-thirds of food and agricultural exports go to OECD countries. Although cereal and vegetable output dropped by 40% between 1990 and 2008, output in grains has since increased, and the 2016–2017 season registered the biggest grain output in a decade. Maize, barley, oats and rice are also grown. Quality Oriental tobacco is a significant industrial crop. Bulgaria is also the largest producer globally of lavender and rose oil, both widely used in fragrances. Within the services sector, tourism is a significant contributor to economic growth. Sofia, Plovdiv, Veliko Tarnovo, coastal resorts Albena, Golden Sands and Sunny Beach and winter resorts Bansko, Pamporovo and Borovets are some of the locations most visited by tourists. Most visitors are Romanian, Turkish, Greek and German. Tourism is additionally encouraged through the 100 Tourist Sites system.
Science and technology. Spending on research and development amounts to 0.78% of GDP, and the bulk of public R&D funding goes to the Bulgarian Academy of Sciences (BAS). Private businesses accounted for more than 73% of R&D expenditures and employed 42% of Bulgaria's 22,000 researchers in 2015. The same year, Bulgaria ranked 39th out of 50 countries in the Bloomberg Innovation Index, the highest score being in education (24th) and the lowest in value-added manufacturing (48th). Bulgaria was ranked 38th in the Global Innovation Index in 2024. Chronic government underinvestment in research since 1990 has forced many professionals in science and engineering to leave Bulgaria. Despite the lack of funding, research in chemistry, materials science and physics remains strong. Antarctic research is actively carried out through the St. Kliment Ohridski Base on Livingston Island in Western Antarctica. The information and communication technologies (ICT) sector generates three per cent of economic output and employs 40,000 to 51,000 software engineers. Bulgaria was known as a "Communist Silicon Valley" during the Soviet era due to its key role in COMECON computing technology production. A concerted effort by the communist government to teach computing and IT skills in schools also indirectly made Bulgaria a major source of computer viruses in the 1980s and 90s. The country is a regional leader in high performance computing: it operates "Avitohol", the most powerful supercomputer in Southeast Europe, and will host one of the eight petascale EuroHPC supercomputers.
Bulgaria has made numerous contributions to space exploration. These include two scientific satellites, more than 200 payloads and 300 experiments in Earth orbit, as well as two cosmonauts since 1971. Bulgaria was the first country to grow wheat in space with its Svet greenhouses on the Mir space station. It was involved in the development of the Granat gamma-ray observatory and the Vega programme, particularly in modelling trajectories and guidance algorithms for both Vega probes. Bulgarian instruments have been used in the exploration of Mars, including a spectrometer that took the first high quality spectroscopic images of Martian moon Phobos with the Phobos 2 probe. Cosmic radiation en route to and around the planet has been mapped by Liulin-ML dosimeters on the ExoMars TGO. Variants of these instruments have also been fitted on the International Space Station and the Chandrayaan-1 lunar probe. Another lunar mission, SpaceIL's "Beresheet", was also equipped with a Bulgarian-manufactured imaging payload. Bulgaria's first geostationary communications satellite—BulgariaSat-1—was launched by SpaceX in 2017.
Infrastructure. Telephone services are widely available, and a central digital trunk line connects most regions. Vivacom (BTC) serves more than 90% of fixed lines and is one of the three operators providing mobile services, along with A1 and Telenor. Internet penetration stood at 69.2% of the population aged 16–74 and 78.9% of households in 2020. Bulgaria's strategic geographic location and well-developed energy sector make it a key European energy centre despite its lack of significant fossil fuel deposits. Thermal power plants generate 48.9% of electricity, followed by nuclear power from the Kozloduy reactors (34.8%) and renewable sources (16.3%). Equipment for a second nuclear power station at Belene has been acquired, but the fate of the project remains uncertain. Installed capacity amounts to 12,668 MW, allowing Bulgaria to exceed domestic demand and export energy. The national road network has a total length of , of which are paved. Railroads are a major mode of freight transportation, although highways carry a progressively larger share of freight. Bulgaria has of railway track, with rail links available to Romania, Turkey, Greece, and Serbia, and express trains serving direct routes to Kyiv, Minsk, Moscow and Saint Petersburg. Sofia is the country's air travel hub, while Varna and Burgas are the principal maritime trade ports.
Demographics. According to the government's official 2022 estimate, the population of Bulgaria consists of 6,447,710 people, down from 6,519,789 according to the last official census in 2021. The majority of the population, 72.5%, reside in urban areas. , Sofia is the most populated urban centre with 1,241,675 people, followed by Plovdiv (346,893), Varna (336,505), Burgas (202,434) and Ruse (142,902). Bulgarians are the main ethnic group and constitute 84.6% of the population. Turkish and Roma minorities account for 8.4 and 4.4%, respectively; some 40 smaller minorities account for 1.3%, and 1.3% do not self-identify with an ethnic group. The Roma minority is usually underestimated in census data and may represent up to 11% of the population. Population density is 55–60 per square kilometre (ultimo 2023), almost half the European Union average. Bulgaria is in a state of demographic crisis. It has had negative population growth since 1989, when the post-Cold War economic collapse caused a long-lasting emigration wave. Some 937,000 to 1,200,000 people—mostly young adults—had left the country by 2005. The majority of children are born to unmarried women. In 2024, the average total fertility rate (TFR) in Bulgaria was 1.59 children per woman, a slight increase from 1.56 in 2018, and well above the all-time low of 1.1 in 1997, but still below the replacement rate of 2.1 and considerably below the historical high of 5.83 children per woman in 1905. Bulgaria thus has one of the oldest populations in the world, with an average age of 43 years. Furthermore, a third of all households consist of only one person and 75.5% of families do not have children under the age of 16. The resulting birth rates are among the lowest in the world while death rates are among the highest.
Bulgaria scores high in gender equality, ranking 18th in the 2018 Global Gender Gap Report. Although women's suffrage was enabled relatively late, in 1937, women today have equal political rights, high workforce participation and legally mandated equal pay. In 2021, market research agency "Reboot Online" ranked Bulgaria as the best European country for women to work. Bulgaria has the highest ratio of female ICT researchers in the EU, as well as the second-highest ratio of females in the technology sector at 44.6% of the workforce. High levels of female participation are a legacy of the Socialist era. Health. High death rates result from a combination of an ageing population, high numbers of people at risk of poverty, and a weak healthcare system. Over 80% of deaths are due to cancer and cardiovascular conditions; nearly a fifth of those are avoidable. Although healthcare in Bulgaria is nominally universal, out-of-pocket expenses account for nearly half of all healthcare spending, significantly limiting access to medical care. Other problems disrupting care provision are the emigration of doctors due to low wages, understaffed and under-equipped regional hospitals, supply shortages and frequent changes to the basic service package for those insured. The 2018 Bloomberg Health Care Efficiency Index ranked Bulgaria last out of 56 countries. Average life expectancy is 74.8 years, compared with an EU average of 80.99 and a world average of 72.38.
Education. Public expenditures for education are far below the European Union average as well. Educational standards were once high, but have declined significantly since the early 2000s. Bulgarian students were among the highest-scoring in the world in terms of reading in 2001, performing better than their Canadian and German counterparts; by 2006, scores in reading, math and science had dropped. By 2018, Programme for International Student Assessment studies found 47% of pupils in the 9th grade to be functionally illiterate in reading and natural sciences. Average basic literacy stands high at 98.4% with no significant difference between sexes. The Ministry of Education and Science partially funds public schools, colleges and universities, sets criteria for textbooks and oversees the publishing process. Education in primary and secondary public schools is free and compulsory. The process spans 12 grades, in which grades one through eight are primary and nine through twelve are secondary level. Higher education consists of a 4-year bachelor degree and a 1-year master's degree. Bulgaria's highest-ranked higher education institution is Sofia University.
Language. Bulgarian is the only language with official status. It belongs to the Slavic group of languages but has a number of grammatical peculiarities that set it apart from other Slavic languages: these include a complex verbal morphology (which also codes for distinctions in evidentiality), the absence of noun cases and infinitives, and the use of a suffixed definite article. Religion. Bulgaria is a secular state with guaranteed freedom of religion by constitution, but Eastern Orthodox Christianity is designated as the traditional religion of the country. Approximately two-thirds of Bulgarians identify as Eastern Orthodox Christians. The Bulgarian Orthodox Church was the first church apart from the Four Ancient Patriarchates of the Eastern Orthodox Church—in Constantinople, Alexandria, Antioch and Jerusalem—and the first national church to gain autocephalous status in 927 AD. The Bulgarian Patriarchate has 12 dioceses and over 2,000 priests. Muslims are the second-largest religious community and constitute approx. 10% of Bulgaria's overall religious makeup. A 2011 survey of 850 Muslims in Bulgaria found 30% self-professing as deeply religious and 50% as just religious. According to the study, some religious teachings, like Islamic funeral, have been traditionally incorporated and are widely practiced while other major ones are less observed, such as the Muslim prayer or abstaining from drinking alcohol, eating pork, and cohabitation.
Other important religions include Roman Catholicism and Judaism, whose history in Bulgaria dates back to the early Middle Ages, the Armenian Apostolic Church, as well as various Protestant denominations, all of which stand for around 2% of Bulgaria's population. An ever increasing number of Bulgarians are either irreligious or unaffiliated with any religion, a percentage that has been growing rapidly over the past 20 years, from 3.9% in 2001, through 9.3% in 2011 and all the way to 15.9% in 2021. According to the most recent census of 2021 the religious denominations of the population are, as follows: Christian (71.5%), Islam (10.8%), other religions (0.1%). Further 12.4% were unaffiliated or did not respond. Culture.
А religious visual arts heritage includes frescoes, murals and icons, many produced by the medieval Tarnovo Artistic School. Like literature, it was not until the National Revival when Bulgarian visual arts began to reemerge. Zahari Zograf was a pioneer of the visual arts in the pre-Liberation era. After the Liberation, Ivan Mrkvička, Anton Mitov, Vladimir Dimitrov, Tsanko Lavrenov and Zlatyu Boyadzhiev introduced newer styles and substance, depicting scenery from Bulgarian villages, old towns and historical subjects. Christo is the most famous Bulgarian artist of the 21st century, known for his outdoor installations. Folk music is by far the most extensive traditional art and has slowly developed throughout the ages as a fusion of Far Eastern, Oriental, medieval Eastern Orthodox and standard Western European tonalities and modes. Bulgarian folk music has a distinctive sound and uses a wide range of traditional instruments, such as gadulka, gaida, kaval and tupan. A distinguishing feature is "extended rhythmical time", which has no equivalent in the rest of European music. The State Television Female Vocal Choir won a Grammy Award in 1990 for its performances of Bulgarian folk music. Written musical composition can be traced back to the works of Yoan Kukuzel (–1360), but modern classical music began with Emanuil Manolov, who composed the first Bulgarian opera in 1890. Pancho Vladigerov and Petko Staynov further enriched symphony, ballet and opera, which singers Ghena Dimitrova, Boris Christoff, Ljuba Welitsch and Nicolai Ghiaurov elevated to a world-class level. Bulgarian performers have gained acclaim in other genres like electropop (Mira Aroyo), jazz (Milcho Leviev) and blends of jazz and folk (Ivo Papazov).
The Bulgarian National Radio, bTV and daily newspapers "Trud", and "24 Chasa" are some of the largest national media outlets. Bulgarian media were described as generally unbiased in their reporting in the early 2000s and print media had no legal restrictions. Since then, freedom of the press has deteriorated to the point where Bulgaria scores 111th globally in the World Press Freedom Index, lower than all European Union members and membership candidate states. The government has diverted EU funds to sympathetic media outlets and bribed others to be less critical on problematic topics, while attacks against individual journalists have increased. Collusion between politicians, oligarchs and the media is widespread. Bulgarian cuisine is similar to that of other Balkan countries and demonstrates strong Turkish and Greek influences. Yogurt, lukanka, banitsa, shopska salad, lyutenitsa and kozunak are among the best-known local foods. Meat consumption is lower than the European average, given a cultural preference for a large variety of salads. Bulgaria was the world's second-largest wine exporter until 1989, but has since lost that position. The 2016 harvest yielded 128 million litres of wine, of which 62 million was exported mainly to Romania, Poland and Russia. Mavrud, Rubin, Shiroka melnishka, Dimiat and Cherven Misket are the typical grapes used in Bulgarian wine. Rakia is a traditional fruit brandy that was consumed in Bulgaria as early as the 14th century.
Sports. Bulgaria appeared at the first modern Olympic games in 1896, when it was represented by Swiss gymnast Charles Champaud. Since then, Bulgarian athletes have won 55 gold, 90 silver, and 85 bronze medals, ranking 25th in the all-time medal table. Weightlifting and wrestling are signature sports of Bulgaria. As of 2025, Bulgaria ranks 2nd in the all-time medal table of the European Weightlifting Championships, 3rd in the all-time medal table of the World Weightlifting Championships, 4th in the all-time medal table for Weightlifting at the Summer Olympics, 3rd in the all-time medal table of the European Wrestling Championships, 6th in the all-time medal table of the World Wrestling Championships and 9th in the all-time medal table for wrestling at the Summer Olympics. Coach Ivan Abadzhiev developed innovative training practices that allowed Bulgaria to dominate weightlifting in the 1980s.Bulgarian athletes have also excelled in boxing, gymnastics, volleyball and tennis. Stefka Kostadinova's world record in women's high jump, achieved at the 1987 World Championships, remained unbroken until 2024, making it one of the longest-standing world records in history. Grigor Dimitrov was the first Bulgarian tennis player in the Top 3 ATP rankings.
Football is the most popular sport in the country. The Bulgarian national football team's best performance was a 4th place finish at the 1994 FIFA World Cup, when the squad was spearheaded by the top goalscorer for the tournament- Hristo Stoichkov. Stoichkov is the most successful Bulgarian player of all time; he was awarded the Golden Boot and the Ballon d'Or and was considered one of the best players in the world while playing for FC Barcelona in the 1990s. CSKA Sofia and Levski Sofia are the most successful clubs domestically and long-standing rivals. Ludogorets is remarkable for having advanced from the local fourth division to the 2014–15 UEFA Champions League group stage in a mere nine years. Placed 39th in 2018, it is Bulgaria's highest-ranked club in UEFA.
Bryozoa 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. 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.
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. 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. 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.
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.
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. 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. 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. "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.
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. 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. 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. The common name "moss animals" is the literal meaning of "Bryozoa", from Greek ('moss') and ('animals'), based on the mossy appearance of encrusting species.
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. 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.
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. 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. 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. 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.
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. 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. There are no nephridia ("little kidneys") or other excretory organs in bryozoa, and it is thought that ammonia diffuses out through the body wall and lophophore. More complex waste products are not excreted but accumulate in the polypide, which degenerates after a few weeks. Some of the old polypide is recycled, but much of it remains as a large mass of dying cells containing accumulated wastes, and this is compressed into a "brown body". When the degeneration is complete, the cystid (outer part of the animal) produces a new polypide, and the brown body remains in the coelom, or in the stomach of the new polypide and is expelled next time the animal defecates.
Respiration and circulation. There are no respiratory organs, heart or blood vessels. Instead, zooids absorb oxygen and eliminate carbon dioxide through diffusion. Bryozoa accomplish diffusion through the use of either a thin membrane (in the case of anascans and some polyzoa) or through pseudopores located on the outer dermis of the zooid. The different bryozoan groups use various methods to share nutrients and oxygen between zooids: some have quite large gaps in the body walls, allowing the coelomic fluid to circulate freely; in others, the funiculi (internal "little ropes") of adjacent zooids connect via small pores in the body wall. Reproduction and life cycles. Zooids of all phylactolaemate species are simultaneous hermaphrodites. Although those of many marine species are protandric, in other words function first as males and then as females, their colonies contain a combination of zooids that are in their male and female stages. In all species the ovaries develop on the inside of the body wall, and the testes on the funiculus connecting the stomach to the body wall. Eggs and sperm are released into the coelom, and sperm exit into the water through pores in the tips of some of the tentacles, and then are captured by the feeding currents of zooids that are producing eggs. Some species' eggs are fertilized externally after being released through a pore between two tentacles, which in some cases is at the tip of a small projection called the "intertentacular organ" in the base of a pair of tentacles. Others' are fertilized internally, in the intertentacular organ or in the coelom.
All phylactolaemates and stenolaemates, and most gymnolaemates, exhibit placentation, and has therefore lecithotrophic (non-feeding) larvae. Except for Cyclostomata and the small gymnolaemate family Epistomiidae, which are viviparous, all are brooders. Phylactolaemata brood their embryos in an internal brood sac, but Gymnolaemata both external membranous sacs, skeletal chambers (ovicells) and internal brooding sacs exist. The developing embryo relies on egg's yolk, extraembryonic nutrition (matrotrophy) or both. In ctenostomes the mother provides a brood chamber for the fertilized eggs, and her polypide disintegrates, providing nourishment to the embryo. Stenolaemates produce specialized zooids to serve as brood chambers, and their eggs divide within this to produce up to 100 identical embryos. Planktotrophic (feeding) larvae are only found in class Gymnolaemata: In the cheilostomatan suborder Malacostegina they are found in the two families Membraniporidae and Electridae, and in the three ctenostome families Alcyonidiidae, Farrellidae, and Hislopiidae. In addition there are a few unconfirmed records, like the solitary form Aethozoid where larvae has never been observed, but which is assumed to have planktotrophic larvae.
The cleavage of bryozoan eggs is biradial, in other words the early stages are bilaterally symmetrical. It is unknown how the coelom forms, since the metamorphosis from larva to adult destroys all of the larva's internal tissues. In many animals the blastopore, an opening in the surface of the early embryo, tunnels through to form the gut. However, in bryozoans the blastopore closes, and a new opening develops to create the mouth. Bryozoan larvae vary in form, but all have a band of cilia round the body which enables them to swim, a tuft of cilia at the top, and an adhesive sac that everts and anchors them when they settle on a surface. Some gymnolaemate species produce cyphonautes larvae which have little yolk but a well-developed mouth and gut, and live as plankton for a considerable time before settling. These larvae have triangular shells of chitin, with one corner at the top and the base open, forming a hood round the downward-facing mouth. In 2006 it was reported that the cilia of cyphonautes larvae use the same range of techniques as those of adults to capture food. Species that brood their embryos form larvae that are nourished by large yolks, have no gut and do not feed, and such larvae quickly settle on a surface. In all marine species the larvae produce cocoons in which they metamorphose completely after settling: the larva's epidermis becomes the lining of the coelom, and the internal tissues are converted to a food reserve that nourishes the developing zooid until it is ready to feed. The larvae of phylactolaemates produce multiple polypides, so that each new colony starts with several zooids. In all species the founder zooids then grow the new colonies by budding clones of themselves. In phylactolaemates, zooids die after producing several clones, so that living zooids are found only round the edges of a colony.
Phylactolaemates can also reproduce asexually by a method that enables a colony's lineage to survive the variable and uncertain conditions of freshwater environments. Throughout summer and autumn they produce disc-shaped statoblasts, masses of cells that function as "survival pods" rather like the gemmules of sponges. Statoblasts form on the funiculus connected to the parent's gut, which nourishes them. As they grow, statoblasts develop protective bivalve-like shells made of chitin. When they mature, some statoblasts stick to the parent colony, some fall to the bottom ("sessoblasts"), some contain air spaces that enable them to float ("floatoblasts"), and some remain in the parent's cystid to re-build the colony if it dies. Statoblasts can remain dormant for considerable periods, and while dormant can survive harsh conditions such as freezing and desiccation. They can be transported across long distances by animals, floating vegetation, currents and winds, and even in the guts of larger animals. When conditions improve, the valves of the shell separate and the cells inside develop into a zooid that tries to form a new colony. "Plumatella emarginata" produces both "sessoblasts", which enable the lineage to control a good territory even if hard times decimate the parent colonies, and "floatoblasts", which spread to new sites. New colonies of "Plumatella repens" produce mainly "sessoblasts" while mature ones switch to "floatoblasts". A study estimated that one group of colonies in a patch measuring produced 800,000 statoblasts.
Cupuladriid Bryozoa are capable of both sexual and asexual reproduction. The sexually reproducing colonies (aclonal) are the result of a larval cupuladriid growing into an adult stage whereas the asexual colonies(clonal) are a result of a fragment of a colony of cupuladriids growing into its own colony. The different forms of reproduction in cupuladriids are achieved through a variety of methods depending on the morphology and classification of the zooid. Ecology. Habitats and distribution. Most marine species live in tropical waters at depths less than . However, a few have been found in deep-sea trenches, especially around cold seeps, and others near the poles. The great majority of bryozoans are sessile. Typically, sessile bryozoans live on hard substrates including rocks, sand or shells. Boring bryozoans leave unique borehole traces after dissolving calcium carbonate substrates. Encrusting forms are much the commonest of these in shallow seas, but erect forms become more common as the depth increases. An example of incrustation on pebbles and cobbles is found in the diverse Pleistocene bryozoans found in northern Japan, where fossils have been found of single stones covered with more than 20 bryozoan species. Sediments with smaller particles, like sand or silt, are usually unsuitable habitat for bryozoans, but tiny colonies have been found encrusting grains of coarse sand. Some bryozoan species specialize in colonizing marine algae, seagrasses, and even mangrove roots; the genus "Amphibiobeania" lives on the leaves of mangrove trees and is called "amphibious" because it can survive regular exposure to air at low tide.
There are a variety of "free-living" bryozoans that live un-attached to a substrate. A few forms such as "Cristatella" can move. Lunulitiform cheilostomes are one group of free-living bryozoans with mobile colonies. They form small round colonies un-attached to any substrate; colonies of the genus Selenaria have been observed to "walk" around using setae. Another cheilostome family, the Cupuladriidae, convergently evolved similarly shaped colonies capable of movement. When observed in an aquarium, Selenaria maculata colonies were recorded to crawl at a speed of one meter per hour, climb over each other, move toward light, and right themselves when turned upside-down. Later study of this genus showed that neuroelectrical activity in the colonies increased in correlation with movement toward light sources. It is theorized that the capacity for movement arose as a side effect when colonies evolved longer setae for unburying themselves from sediment. Other free-living bryozoans are moved freely by waves, currents, or other phenomena. An Antarctic species, "Alcyonidium pelagosphaera", consists of floating colonies. The pelagic species is between in diameter, has the shape of a hollow sphere and consists of a single layer of autozooids. It is still not known if these colonies are pelagic their whole life or only represents a temporarily and previously undescribed juvenile stage. Colonies of the species "Alcyonidium disciforme", which is disc-shaped and similarly free-living, inhabit muddy seabeds in the Arctic and can sequester sand grains they have engulfed, potentially using the sand as ballast to turn themselves right-side-up after they have been overturned. Some bryozoan species can form bryoliths, sphere-shaped free-living colonies that grow outward in all directions as they roll about on the seabed.
In 2014 it was reported that the bryozoan "Fenestrulina rugula" had become a dominant species in parts of Antarctica. Global warming has increased the rate of scouring by icebergs, and this species is particularly adept at recolonizing scoured areas. The phylactolaemates live in all types of freshwater environment – lakes and ponds, rivers and streams, and estuaries – and are among the most abundant sessile freshwater animals. Some ctenostomes are exclusively freshwater while others prefer brackish water but can survive in freshwater. Scientists' knowledge of freshwater bryozoan populations in many parts of the world is incomplete, even in some parts of Europe. It was long thought that some freshwater species occurred worldwide, but since 2002 all of these have been split into more localized species. Bryozoans grow in clonal colonies. A larval Bryozoan settles on a hard substance and produces a colony asexually through budding. These colonies can grow thousands of individual zooids in a relatively short period of time. Even though colonies of zooids grow through asexual reproduction, Bryozoans are hermaphrodites and new colonies can be formed through sexual reproduction and the generation of free swimming larvae. When colonies grow too large, however, they can split in two. This is the only case where asexual reproduction results in a new colony separate from its predecessor. Most colonies are stationary. Indeed, these colonies tend to be settled on immobile substances such as sediment and coarse substances. There are some colonies of freshwater species such as "Cristatella mucedo" that are able to move slowly on a creeping foot.
Interactions with non-human organisms. Marine species are common on coral reefs, but seldom a significant proportion of the total biomass. In temperate waters, the skeletons of dead colonies form a significant component of shell gravels, and live ones are abundant in these areas. The marine lace-like bryozoan "Membranipora membranacea" produces spines in response to predation by several species of sea slugs (nudibranchs). Other predators on marine bryozoans include fish, sea urchins, pycnogonids, crustaceans, mites and starfish. In general marine echinoderms and molluscs eat masses of zooids by gouging pieces of colonies, breaking their mineralized "houses", while most arthropod predators on bryozoans eat individual zooids. In freshwater, bryozoans are among the most important filter feeders, along with sponges and mussels. Freshwater bryozoans are attacked by many predators, including snails, insects, and fish. In Thailand the introduced species "Pomacea canaliculata" (golden apple snail), which is generally a destructive herbivore, has wiped out phylactolaemate populations wherever it has appeared. "P. canaliculata" also preys on a common freshwater gymnolaemate, but with less devastating effect. Indigenous snails do not feed on bryozoans.
Several species of the hydroid family Zancleidae have symbiotic relationships with bryozoans, some of which are beneficial to the hydroids while others are parasitic. Modifications appear in the shapes of some these hydroids, for example smaller tentacles or encrustation of the roots by bryozoans. The bryozoan "Alcyonidium nodosum" protects the whelk "Burnupena papyracea" against predation by the powerful and voracious rock lobster "Jasus lalandii". While whelk shells encrusted by the bryozoans are stronger than those without this reinforcement, chemical defenses produced by the bryozoans are probably the more significant deterrent. In the Banc d'Arguin offshore Mauritania the species "Acanthodesia commensale", which is generally growing attached to gravel and hard-substrate, has formed a facultative symbiotic relationship with hermit crabs of the species "Pseudopagurus cf. granulimanus" resulting in egg-size structures known as bryoliths. Nucleating on an empty gastropod shell, the bryozoan colonies form multilamellar skeletal crusts that produce spherical encrustations and extend the living chamber of the hermit crab through helicospiral tubular growth.
Some phylactolaemate species are intermediate hosts for a group of myxozoa that have also been found to cause proliferative kidney disease, which is often fatal in salmonid fish, and has severely reduced wild fish populations in Europe and North America. "Membranipora membranacea", whose colonies feed and grow exceptionally fast in a wide range of current speeds, was first noticed in the Gulf of Maine in 1987 and quickly became the most abundant organism living on kelps. This invasion reduced the kelp population by breaking their fronds, so that its place as the dominant "vegetation" in some areas was taken by another invader, the large alga "Codium fragile tomentosoides". These changes reduced the area of habitat available for local fish and invertebrates. "M. membranacea" has also invaded the northwest coast of the US. A few freshwater species have been also found thousands of kilometers from their native ranges. Some may have been transported naturally as statoblasts. Others more probably were spread by humans, for example on imported water plants or as stowaways on ships.
Interaction with humans. Fish farms and hatcheries have lost stock to proliferative kidney disease, which is caused by one or more myxozoans that use bryozoans as alternate hosts. Some fishermen in the North Sea have had to find other work because of a form of eczema (a skin disease) known as "Dogger Bank itch", caused by contact with bryozoans that have stuck to nets and lobster pots. Marine bryozoans are often responsible for biofouling on ships' hulls, on docks and marinas, and on offshore structures. They are among the first colonizers of new or recently cleaned structures. Freshwater species are occasional nuisances in water pipes, drinking water purification equipment, sewage treatment facilities, and the cooling pipes of power stations. A group of chemicals called bryostatins can be extracted from the marine bryozoan "Bugula neritina". In 2001 pharmaceutical company GPC Biotech licensed bryostatin 1 from Arizona State University for commercial development as a treatment for cancer. GPC Biotech canceled development in 2003, saying that bryostatin 1 showed little effectiveness and some toxic side effects. In January 2008 a clinical trial was submitted to the United States National Institutes of Health to measure the safety and effectiveness of Bryostatin 1 in the treatment of Alzheimer's disease. However, no participants had been recruited by the end of December 2008, when the study was scheduled for completion. More recent work shows it has positive effects on cognition in patients with Alzheimer's disease with few side effects. About of bryozoans must be processed to extract of bryostatin, As a result, synthetic equivalents have been developed that are simpler to produce and apparently at least as effective.
Biennial plant A biennial plant is a flowering plant that, generally in a temperate climate, takes two years to complete its biological life cycle. Background. In its first year, the biennial plant undergoes primary growth, during which its vegetative structures (leaves, stems, and roots) develop. Usually, the stem of the plant remains short and the leaves are low to the ground, forming a rosette. After one year's growing season, the plant enters a period of dormancy for the colder months. Many biennials require a cold treatment, or vernalization before they will flower. During the next spring or summer, the stem of the biennial plant elongates greatly, or "bolts". The plant then flowers, producing fruits and seeds before it finally dies. There are far fewer biennials than either perennial plants or annual plants. Biennials do not always follow a strict two-year life cycle: most plants in the wild can take three or more years to mature. Rosette leaf size has been found to predict when a plant may enter its second stage of flowering and seed production. Alternatively, under extreme climatic conditions, a biennial plant may complete its life cycle rapidly (e.g., in three months instead of two years). This is quite common in vegetable or flower seedlings that were vernalized before they were planted in the ground. This behavior leads to many normally biennial plants being treated as annuals in some areas. Conversely, an annual grown under extremely favorable conditions may have highly successful seed propagation, giving it the appearance of being biennial or perennial. Some short-lived perennials may appear to be biennial rather than perennial. True biennials flower only once, while many perennials will flower every year once mature.
Biennials grown for flowers, fruits, or seeds are grown for two years, whereas those grown for edible leaves or roots are harvested after one year—and are not kept a second year to run to seed. Examples of biennial plants are members of the onion family including leek, some members of the cabbage family, common mullein, parsley, fennel, "Lunaria", silverbeet, black-eyed Susan, sweet William, colic weed, carrot, and some hollyhocks. Plant breeders have produced annual cultivars of several biennials that will flower the first year from seed, for example, foxglove and stock.
Bay leaf The bay leaf is an aromatic leaf commonly used as a herb in cooking. It can be used whole, either dried or fresh, in which case it is removed from the dish before consumption, or less commonly used in ground form. The flavour that a bay leaf imparts to a dish has not been universally agreed upon, but many agree it is a subtle addition. Bay leaves come from various plants and are used for their distinctive flavour and fragrance. The most common source is the bay laurel ("Laurus nobilis"). Other types include California bay laurel, Indian bay leaf, West Indian bay laurel, and Mexican bay laurel. Bay leaves contain essential oils, such as eucalyptol, terpenes, and methyleugenol, which contribute to their taste and aroma. Bay leaves are used in cuisines including Indian, Filipino, European, and Caribbean. They are typically used in soups, stews, meat, seafood, and vegetable dishes. The leaves should be removed from the cooked food before eating as they can be abrasive in the digestive tract. Bay leaves are used as an insect repellent in pantries and as an active ingredient in killing jars for entomology. In Eastern Orthodoxy liturgy, they are used to symbolize Jesus' destruction of Hades and freeing of the dead.
While some visually similar plants have poisonous leaves, bay leaves are not toxic. However, they remain stiff even after cooking and may pose a choking hazard or cause harm to the digestive tract if swallowed whole or in large pieces. Canadian food and drug regulations set specific standards for bay leaves, including limits on ash content, moisture levels, and essential oil content. Sources. Bay leaves come from several plants, such as: Chemical constituents. The leaves of the European / Mediterranean plant "Laurus nobilis" contain about 1.3% essential oils ("ol. lauri folii"), consisting of 45% eucalyptol, 12% other terpenes, 8–12% terpinyl acetate, 3–4% sesquiterpenes, 3% methyleugenol, and other α- and β-pinenes, phellandrene, linalool, geraniol, terpineol, and also contain lauric acid. Taste and aroma. If eaten whole, "Laurus nobilis" bay leaves are pungent and have a sharp, bitter taste. As with many spices and flavourings, the fragrance of the bay leaf is more noticeable than its taste. When the leaf is dried, the aroma is herbal, slightly floral, and somewhat similar to oregano and thyme. Myrcene, a component of many essential oils used in perfumery, can be extracted from this bay leaf. They also contain eugenol.
Uses. Culinary. Bay leaf is typically used in cooking to flavor broths, grains, soups, stews and stocks. It is typically removed before serving. In Indian cuisine, bay laurel leaves are sometimes used in place of Indian bay leaf, although they have a different flavour. They are most often used in rice dishes like biryani and as an ingredient in garam masala. Bay leaves are called (, in Hindi), Tejpātā (তেজপাতা) in Bengali, তেজ পাত in Assamese and usually rendered into English as Tej Patta. In the Philippines, dried bay laurel leaves are used in several Filipino dishes, such as menudo, beef pares, and adobo. Bay leaves were used for flavouring by the ancient Greeks. They are a fixture in the cooking of many European cuisines (particularly those of the Mediterranean), as well as in the Americas. They are used in soups, stews, brines, meat, seafood, vegetable dishes, and sauces. The leaves also flavour many classic French and Italian dishes. The leaves are most often used whole (sometimes in a ) and removed before serving (they can be abrasive in the digestive tract). Thai and Laotian cuisine employs bay leaf (, ) in a few Arab-influenced dishes, notably massaman curry.
Bay leaves can also be crushed or ground before cooking. Crushed bay leaves impart more fragrance than whole leaves, but are more difficult to remove and thus they are often used in a muslin bag or tea infuser. Ground bay laurel may be substituted for whole leaves and does not need to be removed, but it is much stronger. To brew tea, bay leaves are best boiled for a brief period—typically 3 minutes—to prevent bitterness, as prolonged boiling may overpower the tea's flavour. Fresh bay leaves impart a stronger aroma, while dried leaves require longer steeping for a similar effect. Bay leaves are also used in the making of jerk chicken in the Caribbean Islands. The bay leaves are soaked and placed on the cool side of the grill. Pimento sticks are placed on top of the leaves, and the chicken is placed on top and smoked. The leaves are also added whole to soups, stews, and other Caribbean dishes. Other. Bay leaves can also be used scattered in a pantry to repel meal moths, flies, and cockroaches. Mediouni-Ben Jemaa and Tersim 2011 find the essential oil to be usable as an insect repellent.
Bay leaves have been used in entomology as the active ingredient in killing jars. The crushed, fresh, young leaves are put into the jar under a layer of paper. The vapors they release kill insects slowly but effectively and keep the specimens relaxed and easy to mount. The leaves discourage the growth of molds. They are not effective for killing large beetles and similar specimens, but insects that have been killed in a cyanide killing jar can be transferred to a laurel jar to await mounting. There is confusion in the literature about whether "Laurus nobilis" is a source of cyanide to any practical extent, but there is no evidence that cyanide is relevant to its value in killing jars. It certainly is rich in various essential oil components that could incapacitate insects in high concentrations; such compounds include 1,8-cineole, alpha-terpinyl acetate, and methyl eugenol. It also is unclear to what extent the alleged effect of cyanide released by the crushed leaves has been mis-attributed to "Laurus nobilis" in confusion with the unrelated "Prunus laurocerasus", the so-called cherry laurel, which certainly does contain dangerous concentrations of cyanogenic glycosides together with the enzymes to generate the hydrogen cyanide from the glycocides if the leaf is physically damaged.
Bay leaves are used in Eastern Orthodoxy liturgy. To mark Jesus' destruction of Hades and freeing of the dead, parishioners throw bay leaves and flowers into the air, letting them flutter to the ground. Safety. Some members of the laurel family, as well as the unrelated but visually similar mountain laurel and cherry laurel, have leaves that are poisonous to humans and livestock. While these plants are not sold anywhere for culinary use, their visual similarity to bay leaves has led to the oft-repeated belief that bay leaves should be removed from food after cooking because they are poisonous. This is not true; bay leaves may be eaten without toxic effect. However, they remain unpleasantly stiff even after thorough cooking, and if swallowed whole or in large pieces they may pose a risk of harming the digestive tract or causing choking. Thus, most recipes that use bay leaves will recommend their removal after the cooking process has finished. Canadian food and drug regulations. The Canadian government requires that ground bay leaves contain no more than 4.5% total ash material, with a maximum of 0.5% of which is insoluble in hydrochloric acid. To be considered dried, they must contain 7% moisture or less. The oil content cannot be less than 1 millilitre per 100 grams of the spice.
Basis Basis is a term used in mathematics, finance, science, and other contexts to refer to foundational concepts, valuation measures, or organizational names; here, it may refer to:
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fossil beds containing soft-part imprints. The rock unit is a black shale and crops out at a number of localities near the town of Field in Yoho National Park and the Kicking Horse Pass. Another outcrop is in Kootenay National Park 42 km to the south. History and significance. The Burgess Shale was discovered by palaeontologist Charles Walcott on 30 August 1909, towards the end of the season's fieldwork. He returned in 1910 with his sons, daughter, and wife, establishing a quarry on the flanks of Fossil Ridge. The significance of soft-bodied preservation, and the range of organisms he recognised as new to science, led him to return to the quarry almost every year until 1924. At that point, aged 74, he had amassed over 65,000 specimens. Describing the fossils was a vast task, pursued by Walcott until his death in 1927. Walcott, led by scientific opinion at the time, attempted to categorise all fossils into living taxa, and as a result, the fossils were regarded as little more than curiosities at the time. It was not until 1962 that a first-hand reinvestigation of the fossils was attempted, by Alberto Simonetta. This led scientists to recognise that Walcott had barely scratched the surface of information available in the Burgess Shale, and also made it clear that the organisms did not fit comfortably into modern groups.
Excavations were resumed at the Walcott Quarry by the Geological Survey of Canada under the persuasion of trilobite expert Harry Blackmore Whittington, and a new quarry, the Raymond, was established about 20 metres higher up Fossil Ridge. Whittington, with the help of research students Derek Briggs and Simon Conway Morris of the University of Cambridge, began a thorough reassessment of the Burgess Shale, and revealed that the fauna represented were much more diverse and unusual than Walcott had recognized. Many of the animals present had bizarre anatomical features and only the slightest resemblance to other known animals. Examples include "Opabinia", with five eyes and a snout like a vacuum cleaner hose and "Hallucigenia", which was originally reconstructed upside down, walking on bilaterally symmetrical spines. With Parks Canada and UNESCO recognising the significance of the Burgess Shale, collecting fossils became politically more difficult from the mid-1970s. Collections continued to be made by the Royal Ontario Museum. The curator of invertebrate palaeontology, Desmond Collins, identified a number of additional outcrops, stratigraphically both higher and lower than the original Walcott quarry. These localities continue to yield new organisms faster than they can be studied.
Stephen Jay Gould's book "Wonderful Life", published in 1989, brought the Burgess Shale fossils to the public's attention. Gould suggests that the extraordinary diversity of the fossils indicates that life forms at the time were much more disparate in body form than those that survive today, and that many of the unique lineages were evolutionary experiments that became extinct. Gould's interpretation of the diversity of Cambrian fauna relied heavily on Simon Conway Morris's reinterpretation of Charles Walcott's original publications. However, Conway Morris strongly disagreed with Gould's conclusions, arguing that almost all the Cambrian fauna could be classified into modern day phyla. The Burgess Shale has attracted the interest of paleoclimatologists who want to study and predict long-term future changes in Earth's climate. According to Peter Ward and Donald Brownlee in the 2003 book "The Life and Death of Planet Earth", climatologists study the fossil records in the Burgess Shale to understand the climate of the Cambrian explosion. It can be used to predict what Earth's climate would look like 500 million years in the future as a warming and expanding Sun, combined with declining CO2 and oxygen levels, eventually heat the Earth toward temperatures not seen since the Archean Eon 3 billion years ago (before the first plants and animals appeared). This in turn furthers understanding of how and when the last living things on Earth could potentially die out. See also Future of the Earth.
After the Burgess Shale site was registered as a World Heritage Site in 1980, it was included in the Canadian Rocky Mountain Parks WHS designation in 1984. In 2012, the discovery was announced of another Burgess Shale outcrop in Kootenay National Park to the south. In just 15 days of field collecting in 2013, 50 animal species were unearthed at the new site. IUGS geological heritage site. In respect of the site being "characterized by exceptional soft-tissue preservation, [and containing] the most complete fossil record of Cambrian (Wuliuan) marine ecosystems", the International Union of Geological Sciences (IUGS) included the "Burgess Shale Cambrian Paleontological Record" in its assemblage of 100 "geological heritage sites" around the world in a listing published in October 2022. The organisation defines an "IUGS Geological Heritage Site" as "a key place with geological elements and/or processes of international scientific relevance, used as a reference, and/or with a substantial contribution to the development of geological sciences through history."
Geological setting. The fossil-bearing deposits of the Burgess Shale correlate to the Stephen Formation, a collection of slightly calcareous dark mudstones, about old. The beds were deposited at the base of a cliff about 160 m tall, below the depth agitated by waves during storms. This vertical cliff was composed of the calcareous reefs of the Cathedral Formation, which probably formed shortly before the deposition of the Burgess Shale. The precise formation mechanism is not known for certain, but the most widely accepted hypothesis suggests that the edge of the Cathedral Formation reef became detached from the rest of the reef, slumping and being transported some distance – perhaps kilometers – away from the reef edge. Later reactivation of faults at the base of the formation led to its disintegration from about . This would have left a steep cliff, the bottom of which would be protected from tectonic decompression because the limestone of the Cathedral Formation is difficult to compress. This protection explains why fossils preserved further from the Cathedral Formation are impossible to work with – tectonic squeezing of the beds has produced a vertical cleavage that fractures the rocks, so they split perpendicular to the fossils. The Walcott quarry produced such spectacular fossils because it was so close to the Stephen Formation – indeed the quarry has now been excavated to the very edge of the Cambrian cliff.
It was originally thought that the Burgess Shale was deposited in anoxic conditions, but mounting research shows that oxygen was continually present in the sediment. The anoxic setting had been thought to not only protect the newly dead organisms from decay, but it also created chemical conditions allowing the preservation of the soft parts of the organisms. Further, it reduced the abundance of burrowing organisms – burrows and trackways "are" found in beds containing soft-bodied organisms, but they are rare and generally of limited vertical extent. Brine seeps are an alternative hypothesis; see Burgess Shale type preservation for a more thorough discussion. Stratigraphy. The Burgess Shale Formation comprises 10 members, the most famous being the Walcott Quarry Shale Member comprising the greater phyllopod bed. Taphonomy and diagenesis. There are many other comparable Cambrian "lagerstätten"; indeed such assemblages are far more common in the Cambrian than in any other period. This is mainly due to the limited extent of burrowing activity; as such bioturbation became more prevalent throughout the Cambrian, environments capable of preserving organisms' soft parts became much rarer. (The pre-Cambrian fossil record of animals is sparse and ambiguous, cf ediacaran biota.)
Biota. The biota of the Burgess Shale appears to be typical of middle Cambrian deposits. Although the hard-part bearing organisms make up as little as 14% of the community, these same organisms are found in similar proportions in other Cambrian localities. This means that there is no reason to assume that the organisms without hard parts are exceptional in any way; many appear in other lagerstätten of different age and locations. The biota consists of a range of organisms. Free-swimming (nectonic) organisms are relatively rare, with the majority of organisms being bottom dwelling (benthic) — either moving about (vagrant) or permanently attached to the sea floor (sessile). About two-thirds of the Burgess Shale organisms lived by feeding on the organic content in the muddy sea floor, while almost a third filtered out fine particles from the water column. Under 10% of organisms were predators or scavengers, although since these organisms were larger, the biomass was split equally among each of the filter feeding, deposit feeding, predatory and scavenging organisms.
Many Burgess Shale organisms represent stem group members of the modern animal phyla, though crown group representatives of certain phyla are also present. Working with the Burgess Shale. The fossils of the Burgess Shale are preserved as black carbon films on black shales, and so are difficult to photograph; however, various photographic techniques can improve the quality of the images that can be acquired. Other techniques include backscatter SEM, elemental mapping and "camera lucida" drawing. Once images have been acquired, the effects of decay and taphonomy must be accounted for before a correct anatomical reconstruction can be made. A consideration of the combination of characters allows researchers to establish the taxonomic affinity.
Beavis and Butt-Head Beavis and Butt-Head is an American adult animated sitcom created by Mike Judge. The series follows Beavis and Butt-Head, both voiced by Judge, a pair of teenage slackers characterized by their apathy, lack of intelligence, lowbrow humor and love for hard rock and heavy metal. The original series juxtaposes slice-of-life short subjects—in which the teens embark on low-minded misadventures in their fictional town of Highland, Texas—with the pair watching and commenting on music videos. Judge developed the pair when financing and making his own animated shorts; two of these films, including "Frog Baseball", were broadcast by MTV's animation showcase "Liquid Television". The network commissioned a full series, which over its seven seasons became its most popular program. The original series ended in 1997, but has been twice rebooted, first in 2011 for MTV, and again in 2022 for Paramount+. Starting in 2025, new episodes will air on Comedy Central. During its initial run, "Beavis and Butt-Head" received critical acclaim for its satirical, scathing commentary on society, as well as criticism for its alleged influence on adolescents. The characters became pop culture icons among Gen. X viewers, with their sniggering and dialogue becoming catchphrases. The series was adapted into a theatrical film, the commercially successful "Beavis and Butt-Head Do America" (1996), as well as a sequel, "Beavis and Butt-Head Do the Universe" (2022). The franchise also spawned a comic book series, video games, books, an album, and more.
Premise. Beavis and Butt-Head are unintelligent teenage boys who live in the fictional town of Highland, Texas. "Rolling Stone" described them as "thunderously stupid and excruciatingly ugly". They spend time watching television, drinking unhealthy beverages, eating, and embarking on "mundane, sordid" adventures, which often involve vandalism, abuse, violence, or animal cruelty. According to "The Baltimore Sun", Beavis and Butt-Head are "at their most incorrect when it comes to sexuality and matters of gender. The nicest thing you can say about them in this regard is that they are budding misogynists." When Beavis consumes too much caffeine or sugar, he becomes Cornholio, a hyperactive alter ego. Over the course of the series, Beavis and Butt-Head developed more distinct personalities; Beavis was more hyperactive and significantly less intelligent than his best-friend Butt-Head, who was often more relaxed and tended to be the "brains", what little there were, of the duo's shenanigans. Most episodes integrate sequences where Beavis and Butt-Head watch music videos and offer commentary. They prefer videos with "explosions, loud guitars, screaming and death", and favor rock bands such as the Butthole Surfers, Corrosion of Conformity, and Metallica. Judge said he saw Beavis and Butt-Head as "pretty positive characters, generally speaking ... They usually think everything's pretty cool. Or, in one way or another, everything sucks." He said his perception of the characters changed over the years: "When I first started out with the first show, which was "Frog Baseball", they were just two guys that I would definitely want to keep my distance from ... But, by the end of the series, I would think that two guys like that would at least be fun to sit and watch TV with." Judge composed the series' theme song, which is descended from AC/DC's "Gone Shootin'". Judge later claimed that the central guitar riff of the theme is that song played backwards.
Development. "Beavis and Butt-Head" was created by the American animator Mike Judge. He graduated with a degree in physics but struggled to connect with his work in computer science. In the late 1980s, he began making short animated films on his own; he taught himself how to draw and animate and would shoot his projects with a cheap Bolex 16mm film camera. He made several shorts, including "Frog Baseball", which marked the first appearances of the characters. Judge cold-called networks to pitch this concept, and would send out VHS tapes with prints of his films. The voice of Beavis was based on a kid in his high school calculus class, who would always snicker in a distinct way at their attractive female teacher. The art style of what became "Beavis and Butt-Head" was intentionally disruptive and ugly; Judge wanted it to look like "it was drawn by an insane person." The comic strip "Peanuts" was an unlikely influence: Judge stated that Schulz's line work and sketchy sensibility worked its way into his as well. He was also inspired by the work of John Kricfalusi, and fellow Texas animator Wes Archer and his film "Jack Mack and Rad Boy Go!" Aesthetically, Judge likened the program's best episodes to comfort food: "I think there’s something kind of relaxing about it," he noted. He claimed the wacky comedy of "The Beverly Hillbillies" was an influence on the show. Other elements of the setting are left up to the viewer's imagination: there is little said of the characters' backstories, or their parents, and it's unclear whose house the characters are couch-surfing. This aspect of the show was also inspired by "Peanuts", where the characters also seem to inhabit a liminal world without parents.
MTV bought "Frog Baseball" and two other films to air as part of its late-night animation showcase, "Liquid Television", from which it commissioned the series. Production. 1993–1997: First seven seasons and first film. In September 1992, MTV flew Judge to their New York headquarters to commission a full series of the concept. Executives initially approved 35 episodes; the show's seven-figure budget floored Judge, who had only made the films on his own for $800. The original series aired from 1993 to 1997. The show hired four staff writers, who would work up a treatment to present to Judge, who would then revise it to capture his artistic voice. One early writer, David Felton, found it best to write the characters from a primitive state of mind: "You go to that place in your mind where thoughts begin and then just stay there." The music video commentary was more-or-less improvised by Judge, who recorded them alongside an engineer in Austin. Judge had also tried to have the duo comment on the network's other programs, like "The Real World", but found that recording dialogue on top of dialogue too confusing for viewers.
After writing, the episodes were storyboarded and went to the layout phase of animation, before being transferred to cels and photographed. Judge estimated it could "take anywhere from five months to a year before it's on the air." To save time, the animators made a stock selection of different movements like head turns to simplify editing the program. Though many early seasons were made at MTV's midtown Manhattan campus, Judge preferred to produce the show from his Austin home; in a "Los Angeles Times" piece from 1994, it observes: "Judge makes occasional trips to New York to approve the music videos that will be used in the series and to take care of other business, but generally works by fax, FedEx and video conferencing from Austin." Animation was also partially handled by studios in Korea. The show's first season was animated by J. J. Sedelmaier Productions, while the rest was handled by an in-house animation unit for MTV. Initially, the show's animators condensed the show's art style down to the industry standard of limited animation, which Judge likened to a Saturday morning-type style. He was specific about the show looking intentionally off-kilter: "there's something kind of interesting about seeing drawings animated that look like they were done by a 15-year old in his notebook," he later said. In describing the show's style, Elizabeth Kolbert from "The New York Times" wrote: "They are drawn with purposeful crudeness and their motions have the jerky, seasick quality of marionettes."
Getting the show to a place to where Judge felt it was running smoothly was difficult. He was particularly embarrassed by the first five episodes of the show, with its crude animation style, and was ready to end the show after the second season, when he felt like he was running out of ideas. He claimed he got a "second wind" in the series' third season, which is where he felt like it hit its stride, and he also felt particularly inspired during the series' fifth season. The relentless pace in making the show was wearing him down, which is why he chose to end the series after its 1996 film adaption. 2011: Eighth season. Judge returned to the characters to develop an additional season of the program, which aired in 2011. John Altschuler, formerly a writer for "King of the Hill", told a "Rolling Stone" reporter that he saw signs that Mike Judge was thinking of reviving "Beavis and Butt-Head". On more than one occasion, Judge told the writers that one of their ideas for an episode of "King of the Hill" would work well for "Beavis and Butt-Head";
Judge told the writers that one of their ideas for an episode of "King of the Hill" would work well for "Beavis and Butt-Head"; eventually he concluded, "Maybe we should just actually make some good "Beavis and Butt-Head" episodes." Later, a Lady Gaga video convinced Van Toffler of the tenability of a "Beavis and Butt-Head" revival: "I felt like there was a whole crop of new artists—and what the world sorely missed was the point of view that only Beavis and Butt-Head could bring." As in the old series, Beavis and Butt-Head are high school students who, among other things, criticize contemporary music videos. In updating the show for its millennial-era audience, the duo also watch episodes of "Jersey Shore", Ultimate Fighting Championship matches, and amateur videos from YouTube. Some characters like Daria Morgendorffer did not return. According to TMZ, MTV had not asked Tracy Grandstaff to reprise that role; Judge confirmed that the character was limited to a cameo appearance. As the animation business had switched to digital tools in the time between, this version utilized digital ink and paint, with each drawing simply scanned into a computer and colored.
The episodes debuted in the United States and Canada on October 27, 2011; further, its premiere was a ratings hit, with an audience of 3.3 million total viewers. This number eventually dwindled to 900,000 by the season's end, mainly due to its challenging time slot pitted against regular prime time shows on other networks. As part of a promotional campaign, cinemas screening "Jackass 3D" opened the feature film with a 3-D "Beavis and Butt-Head" short subject. The revival encompassed 24 episodes (12 half-hour programs), but only lasted one season. In a later interview, Judge confirmed "Things didn’t quite click. There were a lot of problems on the animation side of things, trying to get it to look right. I don’t know if that played into it. But it actually did pretty well — the ratings were good. MTV just didn’t want to do anymore." The network's demographic had also shifted to include more female viewers, complicating the show's appeal. In an interview with Howard Stern in 2014, Judge mentioned that while the show's ratings were high, meeting this key demographic was a factor in its cancellation. He also said that MTV was close to selling it to another network, but it became "lost in deal stuff". Judge remained outwardly open to producing more shows, suggesting in 2014 that there was a chance of pitching "Beavis and Butt-Head" to another network. Conflicting with the actual season number, MTV incorrectly refers to this season as "Season 9", even though it is technically the eighth season. Later reruns of this iteration of the program were broadcast on MTV Classic in the mid-2010s. 2022–present: Second film and revival.
Over a decade after the last iteration of the series, the series was again revived, this time in the streaming era for Paramount+. The concept of relaunching the show a second time came from Judge, who created a concert intro for the band Portugal. The Man using the characters. He had not intended to return to the characters again, but found performing the voices fun. He entered discussions with Paramount Global, which was met with more "enthusiasm" than its previous MTV incarnation. The show was originally set to debut on Comedy Central, but was moved to help launch Paramount+. This revival also encompasses additional spin-offs and specials; a second feature film entitled "Beavis and Butt-Head Do the Universe" premiered on June 23, 2022, to kick-off the new series. The new season followed on August 4, 2022, with its second season debuting on April 20, 2023. In addition, in 2022 Paramount+ announced they were to host the full library of over 227 original episodes, newly remastered, with music videos intact for the first time. As of 2024, only some of these episodes have been hosted.
In the new series, Beavis and Butt-Head enter a "whole new Gen Z world" with meta-themes that are said to be relatable to both new fans, who may be unfamiliar with the original series, and old. This iteration has several episodes that depict the characters settling into middle-age, which is a concept Judge had suggested might be amusing over the years. While promoting his film "Extract" in 2009, Judge noted: "I wouldn't mind doing something with them as these two dirty old men sitting on the couch." Season 9 continues the concept of the Beavis and Butt-Head multiverse initially explored in its film predecessor; Teenage Beavis and Butt-Head and Smart Beavis and Butt-Head all get their own dedicated episodes in the revival as well. To promote the revival, Paramount+ attempted to break the world record for the largest serving of nachos at S. Alameda St in Los Angeles to celebrate the return of the show. They were successful and were given a ceremonial plaque from the Guinness World Records representative which stated "The largest serving of nachos was achieved by Mike Judge's Beavis and Butt-Head and Paramount+".
On June 5, 2024, it was announced that the revival had been renewed for a third season, which is expected to air sometime in 2025 on Comedy Central. Live-action adaptation. The concept of taking the teens to the silver screen has been floated since the program was taken on the air. Paramount approached Judge at the start of the show's run to produce a live-action adaption; David Geffen reportedly wanted to cast unknown actors for the role, but Judge refused. Judge has also suggested on more than one occasion directing a live-action adaption of the program. He revealed that Johnny Depp had once expressed interest in the role of Beavis, having imitated the character while Marlon Brando imitated Butt-Head during the production of "Don Juan DeMarco" (1995). He initially disliked the idea of bringing the characters to the real world, but by 2008 had come to believe that "maybe there's something there"; around the same time, he also suggested that "Seann William Scott's kinda got Butt-Head eyes." A decade later, Judge told "Radio Times" "maybe it could be a live-action someday", then went on to speculate that Beavis might be homeless by now. In developing the series' second revival for Paramount+ in the 2020s, executives for the streamer had wanted a live-action "Beavis and Butt-Head" movie. Judge held auditions over Zoom for the project. He eventually talked the company into doing an animated movie instead to reestablish the characters first, with a future live-action movie still a possibility. Judge found it hard to replicate the characters' onscreen stupidity: "It was just sort of like watching teenagers imitate Beavis and Butt-Head."
Reception. Original run. During its original run, "Beavis and Butt-Head" was MTV's highest rated show. It was one of the most popular series when it premiered in 1993. In 1993, "Rolling Stone" described "Beavis and Butt-Head" as the "biggest phenomenon on MTV since the heyday of Michael Jackson". In "Time", Kurt Andersen wrote that "Beavis and Butt-Head" "may be the bravest show ever run on national television". In 1997, Judge said the show was "my reaction to the whole fringe aspects of the political correctness movement". Over its run, "Beavis and Butt-Head" received both positive and negative reactions from the public with its combination of lewd humor and implied criticism of society. It became the focus of criticism from some social critics such as Michael Medved, while others such as David Letterman and the "National Review" defended it as a cleverly subversive vehicle for social criticism and a particularly creative and intelligent comedy. Either way, the show captured the attention of many young television viewers and is often considered a classic piece of 1990s youth culture and Generation X. Trey Parker and Matt Stone, creators of "South Park", cite the series as an influence and compared it to the blues.
In 1997, Dan Tobin of "The Boston Phoenix" commented on the series' humor, saying it transformed "stupidity into a crusade, forcing us to acknowledge how little it really takes to make us laugh." In 1997, Ted Drozdowski of "The Boston Phoenix" described the 1997 "Beavis and Butt-Head" state as "reduced to self-parody of their self-parody". In the "Baltimore Sun", David Zurawik said that "Beavis and Butt-Head" was "intelligent social satire that especially speaks in a meaningful way to a generation of teenage boys who are going through a uniquely complicated socialization at the hands of their baby-boomer parents". He said that its popularity may have taught audiences about male adolescence in the 1990s; he wrote that they were the postmodern descendants of Tom Sawyer and Huckleberry Finn, who were the "exemplars of males coming of age in American popular culture". Legacy. In December 2005, "TV Guide" ranked the duo's distinct laughing at #66 on their list of the 100 Greatest TV Quotes and Catchphrases. In 2012, "TV Guide" ranked "Beavis and Butt-Head" as one of the top 60 Greatest TV Cartoons of All Time. IGN ranked the series as the fifth greatest cartoon of all time on their list of the Top 100 Animated Series.
Various television series have done parodies or homages to the series, including "The Simpsons", "Family Guy", "Arthur", "Friends", "The Critic", "Two and a Half Men", "Frasier", "The Fairly OddParents", "Boy Meets World", "Full House", "", "Robot Chicken", "Married... with Children", "Roseanne", "Tiny Toon Adventures", "Big Mouth", "Drawn Together", "Space Ghost Coast to Coast", "Beverly Hills, 90210", "2 Broke Girls", among others. On the April 13, 2024 episode of "Saturday Night Live" guest host Ryan Gosling, and cast regular Mikey Day, appeared in a sketch as live-action versions of Beavis and Butt-Head. The sketch went viral and received large media coverage and critical acclaim. The pair reprised the roles on the red carpet at "The Fall Guy" movie premiere. Controversies. In its heyday, "Beavis and Butt-Head" became a lightning rod for controversy over its content. "The downward spiral of the living white male surely ends here," John Leland wrote in "Newsweek" in 1993. The show was blamed for the death of two-year-old Jessica Matthews in Moraine, Ohio, in October 1993. The girl's five-year-old brother, Austin Messner, set fire to his mother's mobile home with a cigarette lighter, killing the two-year-old. The mother later claimed that her son watched an episode in which the characters said "fire was fun". However, the neighbors stated that the family did not even have cable television and would thus be unable to view the show.
As a result, all references to fire were removed from subsequent airings and prompted the show to a later time slot. The creators found a censorship loophole and took delight in sometimes making Beavis scream things that sounded very similar to his previous "Fire! Fire!" (such as "Fryer! Fryer!" when he and Butt-Head are working the late shift at Burger World) and also having him almost say the forbidden word (such as one time when he sang "Liar, liar, pants on..." and pausing before "fire"). There was also a music video where a man runs on fire in slow motion ("California" by Wax). Beavis is hypnotized by it and can barely say "fire". However, MTV eventually removed the episode entirely, leading it to be locked away in the MTV vault. References to fire were cut from earlier episodes—even the original master tapes were altered permanently. Other episodes MTV opted not to rerun included "Stewart's House" and "Way Down Mexico Way". Copies of early episodes with the controversial content intact are rare, and the copies that exist are made from home video recordings of the original broadcasts, typically on VHS. In an interview included with the , Judge said he is uncertain whether some of the earlier episodes still exist in their original, uncensored form.

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