European bison
The European bison ("Bison bonasus") or the European wood bison, also known as the wisent ( or ), or the zubr (), or colloquially the European buffalo, is a European species of bison. It is one of two extant species of bison, alongside the American bison. The European bison is the heaviest wild land animal in Europe, and individuals in the past may have been even larger than their modern-day descendants. During late antiquity and the Middle Ages, bison became extinct in much of Europe and Asia, surviving into the 20th century only in northern-central Europe and the northern  Caucasus Mountains. During the early years of the 20th century, bison were hunted to extinction in the wild. The species — now numbering several thousand and returned to the wild by captive breeding programmes — is no longer in immediate danger of extinction, but remains absent from most of its historical range. It is not to be confused with the aurochs ("Bos primigenius"), the extinct ancestor of domestic cattle, with which it once co-existed. Besides humans, bison have few predators. In the 19th century, there were scattered reports of wolves, lions, tigers, and bears hunting bison. In the past, especially during  the Middle Ages, humans commonly killed bison for their hide and meat. They used their horns to make drinking horns. European bison were hunted to extinction in the wild in the early 20th century, with the last wild animals of the "B. b. bonasus" subspecies being shot in the Białowieża Forest (on today's Belarus–Poland border) in 1921. The last of the Caucasian wisent subspecies ("B. b. caucasicus") was shot in the north-western Caucasus in 1927. The Carpathian wisent ("B. b. hungarorum") had been hunted to extinction in the mid-1800s. The Białowieża or lowland European bison was kept  alive in captivity, and has since been reintroduced into several countries in Europe. In 1996, the International Union for Conservation of Nature classified the European bison as an endangered species, no longer extinct in the wild. Its status has improved since then, changing to vulnerable and later to near threatened. European bison were first scientifically described by Carl Linnaeus in 1758. Some later descriptions treat the European bison as conspecific with the American bison. Three subspecies of European bison existed in the recent past, but only one, the nominate subspecies ("B. b. bonasus"), survives today. The ancestry and relationships of  the wisent to fossil bison species remain controversial and disputed. The European bison is one of the national animals of Poland and Belarus. Etymology. The ancient Greeks and ancient Romans were the first to name bison as such; the 2nd-century AD authors Pausanias and Oppian referred to them as . Earlier, in the 4th century BC, during the Hellenistic period, Aristotle referred to bison as . He also noted that the Paeonians called it μόναπος ("monapos"). Claudius Aelianus, writing in the late 2nd or early 3rd centuries AD, also referred to the species as , and both Pliny the Elder's "Natural  History" and Gaius Julius Solinus used and . Both Martial and Seneca the Younger mention (. ). Later Latin spellings of the term included , , and . John Trevisa is the earliest author cited by the "Oxford English Dictionary" as using, in his 1398 translation of Bartholomeus Anglicus's "De proprietatibus rerum", the Latin plural in English, as "bysontes" ( and ). Philemon Holland's 1601 translation of Pliny's "Natural History", referred to "bisontes". The marginalia of the King James Version gives "bison" as a gloss for the Biblical animal called the "pygarg" mentioned in the "Book of Deuteronomy". Randle Cotgrave's 1611 French–English dictionary  notes that was already in use, and it may have influenced the adoption of the word into English, or alternatively it may have been borrowed direct from Latin. John Minsheu's 1617 lexicon, "Ductor in linguas", gives a definition for "Bíson" (). In the 18th century the name of the European animal was applied to the closely related American bison (initially in Latin in 1693, by John Ray) and the Indian bison (the gaur, "Bos gaurus"). Historically the word was also applied to Indian domestic cattle, the zebu ("B. indicus" or "B. primigenius indicus"). Because of the scarcity of the European  bison the word ‘bison’ was most familiar in relation to the American species. By the time of the adoption of ‘bison’ into Early Modern English, the early medieval English name for the species had long been obsolete: the had descended from , and was related to . The word ’wisent’ was then borrowed in the 19th century from modern , itself related to , , , and to , , , and ultimately, like the Old English name, from Proto-Germanic. The , also found in "weasel", originally referred to the animal's musk. The word ‘zubr’ in English is a borrowing from , previously also used to denote one race  of the European bison. The Polish "żubr" is similar to the word for the European bison in other modern Slavic languages, such as and . The noun for the European bison in all living Slavonic tongues is thought to be derived from Proto-Slavic: "*zǫbrъ" ~ "*izǫbrъ", which itself possibly comes from Proto-Indo-European: "*ǵómbʰ-" for tooth, horn or peg. Description. The European bison is the heaviest surviving wild land animal in Europe. Similar to their American cousins, European bisons were potentially larger historically than remnant descendants; modern animals are about in length, not counting a tail of , in height , and in weight for males, and about in body length without tails, in height, and in weight for females. At birth, calves are quite small, weighing between . In the free-ranging population of the Białowieża Forest of Belarus and Poland, body masses among adults (aged 6 and over) are on average in the cases of males, and among females. An occasional big bull European bison can weigh up to or more with old bull records of for lowland wisent and for Caucasian wisent. On average, it is lighter in body mass, and yet slightly taller at the shoulder, than its  American relatives, the wood bison ("Bison bison athabascae") and the plains bison ("Bison bison bison"). Compared to the American species, the wisent has shorter hair on the neck, head, and forequarters, but longer tail and horns. See differences from American bison. The European bison makes a variety of vocalisations depending on its mood and behaviour, but when anxious, it emits a growl-like sound, known in Polish as "chruczenie" (). This sound can also be heard from wisent males during the mating season. History. Historically, the lowland European bison's range encompassed most of the lowlands of northern Europe, extending from  the Massif Central to the Volga River and the Caucasus. It may have once lived in the Asiatic part of what is now the Russian Federation. The European bison is known in southern Sweden only between 9500 and 8700 BP, and in Denmark similarly is documented only from the Pre-Boreal. It is not recorded from the British Isles, nor from Italy or the Iberian Peninsula, although prehistorical absence of the species among British Isles is debatable, based on fossils found on Doggerland or Brown Bank, and Isle of Wight and Oxfordshire, followed by fossil records of steppe bison from  the isles. The extinct steppe bison, "B. priscus", is known from across Eurasia and North America, last occurring 7,000 BC to 5,400 BC, and is depicted in the Cave of Altamira and Lascaux. The Pleistocene woodland bison ("B. schoetensacki"), has been proposed to have last existed around 36,000 BC. But other authors restrict "B. schoetensacki" to remains that are hundreds of thousands of years older. Cave paintings appear to distinguish between "B. bonasus" and "B. priscus". Within mainland Europe, its range decreased as human populations expanded and cut down forests. They seemed to be common in Aristotle's period on  Mount Mesapion (possibly the modern Ograzhden). In the same wider area Pausanias calling them "Paeonian bulls" and "bisons", gives details on how they were captured alive; adding also the fact that a golden Paeonian bull head was offered to Delphi by the Paeonian king Dropion (3rd century BC) who lived in what is today Tikveš. The last references (Oppian, Claudius Aelianus) to the animal in the transitional Mediterranean/Continental biogeographical region in the Balkans in the area of modern borderline between Greece, North Macedonia and Bulgaria date to the 3rd century AD. In northern Bulgaria, the wisent survived until the  9th or 10th century AD. There is a possibility that the species' range extended to East Thrace during the 7th – 8th century AD. Its population in Gaul was extinct in the 8th century AD. The species survived in the Ardennes and the Vosges Mountains until the 15th century. In the Early Middle Ages, the wisent apparently still occurred in the forest steppes east of the Urals, in the Altai Mountains, and seems to have reached Lake Baikal in the east. The northern boundary in the Holocene was probably around 60°N in Finland. European bison survived in a few  natural forests in Europe, but their numbers dwindled. In the Polish–Lithuanian Commonwealth, European bison in the Białowieża Forest were legally the property of the Grand Dukes of Lithuania until the third partition of Poland. Wild European bison herds also existed in the forest until the mid-17th century. Polish kings took measures to protect the bison. King Sigismund II Augustus instituted the death penalty for poaching a European bison in Białowieża in the mid-16th century. In the early 19th century, after the partitions of the Polish Commonwealth the Russian tsars retained old Lithuanian laws protecting the European bison herd in  Białowieża. Despite these measures and others, the European bison population continued to decline over the following century, with only Białowieża and Northern Caucasus populations surviving into the 20th century. The last European bison in Transylvania died in 1790. During World War I, occupying German troops killed 600 of the European bison in the Białowieża Forest for sport, meat, hides and horns. A German scientist informed army officers that the European bison were facing imminent extinction, but at the very end of the war, retreating German soldiers shot all but nine animals. The last wild European bison in Poland was killed  in 1921. The last wild European bison in the world was killed by poachers in 1927 in the western Caucasus. By that year, fewer than 50 remained, all held by zoos. The International Society for the Preservation of the Wisent was founded on 25 and 26 August 1923 in Berlin. Numerous experts joined the company, among them Hermann Pohle, Max Hilzheimer and Julius Riemer. The aim of the society was to cooperate internationally to preserve the Wisent, which was directly threatened with extinction. The last free-living wisent in the Caucasus was shot in 1927. The first goal of the  society was to record all the still living bison, on the basis of which one could begin with a conservation breeding. The company, the first chairman of which was the director of the Frankfurt Zoo, Kurt Priemel, was joined by several private individuals and institutions, such as the American Bison Society, following the example of which the International Society for the Conservation of the Wisent was founded. Important members were, however, the Polish Hunting Association and the Poznań zoological gardens, as well as a number of Polish private individuals. They were also the first to provide substantial funds to acquire  the first bison cows and bulls. The breeding book was published in the company's annual report from 1932. While Priemel aimed at a slow increase in the Wisent population with the pure conservation of the breeding line, Lutz Heck planned to suddenly increase the Wisent population by crossing American bison in 1934 in a separate breeding project in Munich. In it he was personally supported by the then Reichsjägermeister Hermann Göring, who hoped for huntable big game. Lutz Heck promised his powerful supporter in writing: "Since surplus bulls will soon be set, the hunting of the Wisent will be  possible again in the foreseeable future." Hermann Göring himself took over the patronage of the "German Professional Association of Wisent Breeders and Hegers", founded at the suggestion of Lutz Heck. Kurt Priemel, who had since resigned as president of the "International Society for the Preservation of the Wisent", warned in vain against a "manification". This criticism led Lutz Heck to threaten Priemel by announcing that Göring would take action against Priemel if he continued to oppose his crossing plans. Priemel was then banned from publishing in relation to bison breeding, and the regular bookkeeper of the "International Society", Erna Mohr , was also forced to hand over the official register in 1937. Thus, the older society was effectively incorporated into the newly created "professional community" and thus connected to the same. After the Second World War, therefore, only the pure-blooded bison in the game park Springe near Hanover were recognized as part of the international herd book. The first two bison were released into nature to the Białowieża Forest in 1952. By 1964 more than 100 existed. Genetic history. A 2003 study of mitochondrial DNA indicated four distinct maternal lineages in the tribe Bovini: Y chromosome analysis associated wisent and  American bison. An earlier study, using amplified fragment-length polymorphism fingerprinting, showed a close association of wisent and American bison and probably with yak. It noted the interbreeding of Bovini species made determining relationships problematic. European bison can crossbreed with American bison. This hybrid is known in Poland as a "". The products of a German interbreeding programme were destroyed after the Second World War. This programme was related to the impulse which created the Heck cattle. The cross-bred individuals created at other zoos were eliminated from breed books by the 1950s. A Russian back-breeding programme resulted in a  wild herd of hybrid animals, which presently lives in the Caucasian Biosphere Reserve (550 animals in 1999). Wisent-cattle hybrids also occur, similarly to the North American beefalo. Cattle and European bison hybridise fairly readily. First-generation males are infertile. In 1847, a herd of wisent-cattle hybrids named "żubroń" () was created by Leopold Walicki. The animals were intended to become durable and cheap alternatives to cattle. The experiment was continued by researchers from the Polish Academy of Sciences until the late 1980s. Although the program resulted in a quite successful animal that was both hardy and could be bred  in marginal grazing lands, it was eventually discontinued. Currently, the only surviving żubroń herd consists of just a few animals in Białowieża Forest, Poland and Belarus. In 2016, the first whole genome sequencing data from two European bison bulls from the Białowieża Forest revealed that the bison and bovine species diverged from about 1.7 to 0.85 Mya, through a speciation process involving limited gene flow. These data further support the occurrence of more recent secondary contacts, posterior to the divergence between "Bos primigenius primigenius" and "B. p. namadicus" (ca. 150,000 years ago), between the wisent and (European) taurine cattle lineages . An independent study of mitochondrial DNA and autosomal markers confirmed these secondary contacts (with an estimate of up to 10% of bovine ancestry in the modern wisent genome) leading the authors to go further in their conclusions by proposing the wisent to be a hybrid between steppe bison and aurochs with a hybridisation event originating before 120,000 years ago. However, other studies considered the 10% estimate for aurochs gene flow a gross overstimate and based on flawed data, and not supported by the data from the full nuclear genome of the wisent. Some of the authors however support the hypothesis  that similarity of wisent and cattle ("Bos") mitochondrial genomes is result of incomplete lineage sorting during divergence of "Bos" and "Bison" from their common ancestors rather than further post-speciation gene flow (ancient hybridisation between "Bos" and "Bison"). But they agree that limited gene flow from "Bos primigenius taurus" could account for the affiliation between wisent and cattle nuclear genomes (in contrast to mitochondrial ones). Alternatively, genome sequencing completed on remains attributed to the Pleistocene woodland bison ("B. schoetensacki"), and published in 2017, posited that genetic similarities between the Pleistocene woodland bison and the wisent suggest that "B. schoetensaki" was  the ancestor of the European wisent. However, other studies have disputed the attribution of the remains to this species, otherwise known from remains hundreds of thousands of years older, instead referring them to an unnamed lineage of bison closely related to "B. bonasius". A 2018 study proposed that the lineage leading to the wisent and to the lineage ancestral to both the American bison and "Bison priscus" had split over 1 million years ago, with the mitochondrial DNA discrepancy being the result of incomplete lineage sorting. The authors also proposed that during the late Middle Pleistocene there had been interbreeding  between the ancestor of the wisent and the common ancestor of "Bison priscus" and the American bison. Behaviour and biology. Social structure and territorial behaviours. The European bison is a herd animal, which lives in both mixed and solely male groups. Mixed groups consist of adult females, calves, young aged 2–3 years, and young adult bulls. The average herd size is dependent on environmental factors, though on average, they number eight to 13 animals per herd. Herds consisting solely of bulls are smaller than mixed ones, containing two individuals on average. European bison herds are not family units. Different herds  frequently interact, combine, and quickly split after exchanging individuals. Bison social structure has been described by specialists as a matriarchy, as it is the cows of the herd that lead it, and decide where the entire group moves to graze. Although larger and heavier than the females, the oldest and most powerful male bulls are usually satellites that hang around the edges of the herd to protect the group. Bulls begin to serve a more active role in the herd when a danger to the group's safety appears, as well as during the mating season – when they compete with  each other. Territory held by bulls is correlated by age, with young bulls aged between five and six tending to form larger home ranges than older males. The European bison does not defend territory, and herd ranges tend to greatly overlap. Core areas of territory are usually sited near meadows and water sources. Reproduction. The rutting season occurs from August through to October. Bulls aged 4–6 years, though sexually mature, are prevented from mating by older bulls. Cows usually have a gestation period of 264 days, and typically give birth to one calf at a time. On average, male calves  weigh at birth, and females . Body size in males increases proportionately to the age of 6 years. While females have a higher increase in body mass in their first year, their growth rate is comparatively slower than that of males by the age of 3–5. Bulls reach sexual maturity at the age of two, while cows do so in their third year. European bison have lived as long as 30 years in captivity, but in the wild their lifespans are shorter. The lifespan of a zubr in the wild is usually between 18 and 24 years, though females live longer  than males. Productive breeding years are between four and 20 years of age in females, and only between six and 12 years of age in males. Diet. European bison feed predominantly on grasses, although they also browse on shoots and leaves; in summer, an adult male can consume 32 kg of food in a day. European bison in the Białowieża Forest in Poland have traditionally been fed hay in the winter for centuries, and large herds may gather around this diet supplement. European bison need to drink every day, and in winter can be seen breaking ice with their heavy  hooves. Despite their usual slow movements, European bison are surprisingly agile and can clear 3-m-wide streams or 2-m-high fences from a standing start. Differences from American bison. Although superficially similar, a number of physical and behavioural differences are seen between the European bison and the American bison. The zubr has 14 pairs of ribs, while the American bison has 15. Adult European bison are (on average) taller than American bison, and have longer legs. European bison tend to browse more, and graze less than their American relatives, due to their necks being set differently. Compared to  the American bison, the nose of the European bison is set further forward than the forehead when the neck is in a neutral position. The body of the wisent is less hairy, though its tail is hairier than that of the American species. The horns of the European bison point forward through the plane of their faces, making them more adept at fighting through the interlocking of horns in the same manner as domestic cattle, unlike the American bison, which favours charging. European bison are less tameable than the American ones, and breed with domestic cattle less readily. In terms  of behavioural capability, European bison runs slower and with less stamina yet jumps higher and longer than American bisons, showing signs of more developed adaptations into mountainous habitats. Conservation. The protection of the European bison has a long history; between the 15th and 18th centuries, those in the forest of Białowieża were protected and their diet supplemented. Efforts to restore this species to the wild began in 1929, with the establishment of the Bison Restitution Centre at Białowieża, Poland. Subsequently, in 1948, the Bison Breeding Centre was established within the Prioksko-Terrasny Biosphere Reserve. The modern herds are managed as  two separate lines – one consisting of only "Bison bonasus bonasus" (all descended from only seven animals) and one consisting of all 12 ancestors, including the one "B. b. caucasicus" bull. The latter is generally not considered a separate subspecies because they contain DNA from both "B. b. bonasus" and "B. b. caucasicius", although some scientists classify them as a new subspecies, "B. b. montanus". Only a limited amount of inbreeding depression from the population bottleneck has been found, having a small effect on skeletal growth in cows and a small rise in calf mortality. Genetic variability continues to shrink. From  five initial bulls, all current European bison bulls have one of only two remaining Y chromosomes. Reintroduction. Beginning in 1951, European bison have been reintroduced into the wild, including some areas where they were never found wild. Free-ranging herds are currently found in Poland, Lithuania, Belarus, Ukraine, Bulgaria, Romania, Russia, Slovakia, Latvia, Switzerland, Kyrgyzstan, Germany, and in forest preserves in the Western Caucasus. The Białowieża Primeval Forest, an ancient woodland that straddles the border between Poland and Belarus, continues to have the largest free-living zubr population in the world with around 1000 wild bison counted in 2014. Herds  have also been introduced in Moldova (2005), Spain (2010), Denmark (2012), and the Czech Republic (2014). The Wilder Blean project, headed up by the Wildwood Trust and Kent Wildlife Trust, is introducing European bison to the UK for the first time in 6000 years. The herd of 3 females and 1 male will be set free in 2022 within a 2,500-acre conservation area in Blean Woods near Canterbury. In March 2022, five animals (one bull and four cows) were reintroduced in Serbia, where bison went extinct c.1800. Animals were transported from the Białowieża Forest and reintroduced on the Fruška  Gora mountain. Numbers and distribution. Numbers by country. The total worldwide population recorded in 2019 was around 7,500 – about half of this number being in Poland and Belarus, with over 25% of the global population based in Poland alone. For 2016, the number was 6,573 (including 4,472 free-ranging) and has been increasing. Some local populations are estimated as: Distribution. The largest zubr herds — of both captive and wild populations — are still based in Poland and Belarus, the majority of which can be found in the Białowieża Forest including the most numerous population of free-living European bison in the  world with most of the animals living on the Polish side of the border. Poland remains the world's breeding centre for the wisent. In the years 1945 to 2014, from the Białowieża National Park alone, 553 specimens were sent to most captive populations of the zubr in Europe as well as all breeding sanctuaries for the species in Poland. Since 1983, a small reintroduced population lives in the Altai Mountains. This population suffers from inbreeding depression and needs the introduction of unrelated animals for "blood refreshment". In the long term, authorities hope to establish a population of about 1,000  animals in the area. One of the northernmost current populations of the European bison lives in Vologodskaya Oblast in the Northern Dvina valley at about 60°N. It survives without supplementary winter feeding. Another Russian population lives in the forests around the Desna River on the border between Russia and Ukraine. The north-easternmost population lives in Pleistocene Park south of Chersky in Siberia, a project to recreate the steppe ecosystem which began to be altered 10,000 years ago. Five wisents were introduced on 24 April 2011. The wisents were brought to the park from the Prioksko-Terrasny Nature  Reserve near Moscow. Winter temperatures often drop below −50 °C. Four of the five bison have subsequently died due to problems acclimatizing to the low winter temperature. In 2011, three bison were introduced into Alladale Wilderness Reserve in Scotland. Plans to move more into the reserve were made, but the project failed due to not being "well thought through". In April 2013, eight European bison (one male, five females, and two calves) were released into the wild in the Bad Berleburg region of Germany, after 850 years of absence since the species became extinct in that region. Plans are  being made to reintroduce two herds in Germany and in the Netherlands in Oostvaardersplassen Nature Reserve in Flevoland as well as the Veluwe. In 2007, a bison pilot project in a fenced area was begun in Zuid-Kennemerland National Park in the Netherlands. Because of their limited genetic pool, they are considered highly vulnerable to illnesses such as foot-and-mouth disease. In March 2016, a herd was released in the Maashorst Nature Reserve in North Brabant. Zoos in 30 countries also have quite a few bison involved in captive-breeding programs. Cultural significance. Representations of the European bison from  different ages, across millennia of human society's existence, can be found throughout Eurasia in the form of drawings and rock carvings; one of the oldest and most famous instances of the latter can be found in the Cave of Altamira, present-day Spain, where cave art featuring the wisent from the Upper Paleolithic was discovered. The bison has also been represented in a wide range of art in human history, such as sculptures, paintings, photographs, glass art, and more. Sculptures of the wisent constructed in the 19th and 20th centuries continue to stand in a number of European cities ; arguably the most notable of these are the designed by Mihály Zichy and the two bison sculptures in Kiel sculpted by August Gaul in 1910–1913. However, a number of other monuments to the animal also exist, such as those in Hajnówka and Pszczyna or at the Kyiv Zoo entrance. Mikołaj Hussowczyk, a poet writing in Latin about the Grand Duchy of Lithuania during the early 16th century, described the zubr in a historically significant fictional work from 1523. The zubr is considered one of the national animals of Poland and Belarus. Due to this and the fact that half of  the worldwide European bison population can be found spread across these two countries, the wisent is still featured prominently in the heraldry of these neighbouring states (especially in the overlapping region of Eastern Poland and Western Belarus). Examples in Poland include the coats of arms of: the counties of Hajnówka and Zambrów, the towns Sokółka and Żywiec, the villages Białowieża and Narewka, as well as the coats of arms of the Pomian and Wieniawa families. Examples in Belarus include the Grodno and Brest voblasts, the town of Svislach, and others. The European bison can also be found on the coats  of arms of places in neighbouring countries: Perloja in southern Lithuania, Lypovets in west-central Ukraine, and Zubří in east Czechia – as well as further outside the region, such as Kortezubi in the Basque Country, and Jabel in Germany. A flavoured vodka called Żubrówka (), originating as a recipe of the szlachta of the Kingdom of Poland in the 14th century, has since 1928 been industrially produced as a brand in Poland. In the decades that followed, it became known as the "world's best known Polish vodka" and sparked the creation of a number of copy brands inspired by the  original in Belarus, Russia, Germany, as well as other brands in Poland. The original Polish brand is known for placing a decorative blade of bison grass from the Białowieża Forest in each bottle of their product; both the plant's name in Polish and the vodka are named after "żubr", the Polish name for the European bison. The zubr also appears commercially as a symbol of a number of other Polish brands, such as the popular beer brand Żubr and on the logo of Poland's second largest bank, Bank Pekao S.A.