Theaurochs (Bos primigenius;/ˈɔːrɒks/ or/ˈaʊrɒks/; pl.:aurochs oraurochsen) is anextinct species ofbovine, considered to be the wild ancestor of modern domesticcattle. With a shoulder height of up to 180 cm (71 in) in bulls and 155 cm (61 in) in cows, it was one of the largestherbivores in theHolocene; it had massive elongated and broad horns that reached 80 cm (31 in) in length.
The aurochs was part of thePleistocene megafauna. It probably evolved in Asia and migrated west and north during warminterglacial periods. The oldest-known aurochs fossils date to theMiddle Pleistocene. The species had an expansive range spanning from Western Europe and North Africa to the Indian subcontinent and East Asia. The distribution of the aurochs progressively contracted during the Holocene due tohabitat loss and hunting, with the last known individual dying in theJaktorów forest in Poland in 1627.
Two aurochsdomestication events occurred during theNeolithic Revolution. One gave rise to the domestictaurine cattle (Bos taurus) in theFertile Crescent in theNear East that was introduced to Europe via theBalkans and the coast of theMediterranean Sea.Hybridisation between aurochs and early domestic cattle occurred during the early Holocene. Domestication of theIndian aurochs led to thezebu cattle (Bos indicus) that hybridised with early taurine cattle in theNear East about 4,000 years ago. Some modern cattle breeds exhibit features reminiscent of the aurochs, such as the dark colour and light eel stripe along the back of bulls, the lighter colour of cows, or an aurochs-like horn shape.
The use of the plural formaurochsen in English is a direct parallel of the German pluralOchsen and recreates the same distinction by analogy as English singularox and pluraloxen, althoughaurochs may stand for both the singular and the plural term; both are attested.[7][8]
The North African aurochs (B. p. mauritanicus) lived north of theSahara.[5] This subspecies has also been calledB. p. opisthonomus.[17]
In the 21st century, Chinese geneticists publishedmitochondrial DNA evidence supporting that Eurasian aurochs populations fromnorthern China were genetically isolated for large stretches of thePleistocene, and as a result distinctive enough to be considered a separate subspecies, the East Asian aurochs (B. p. sinensis), even if the animals were not morphologically distinct.[17]
At least twodwarf subspecies of aurochs developed onMediterranean islands as a result of sea level changes during the Pleistocene:
The coldPliocene climate caused an extension of opengrassland, which enabled theevolution of large grazers.[5] The origin of the aurochs is unclear, with authors suggesting either an African or Asian origin for the species.Bos acutifrons is considered to be a possible ancestor of the aurochs, of which a fossil skull wasexcavated in theSivalik Hills in India that dates to theEarly Pleistocene about2 million years ago.[26]
An aurochs skull excavated in Tunisia'sKef Governorate from earlyMiddle Pleistocenestrata dating about0.78 million years ago is the oldest well-dated fossil specimen to date. The authors of the study proposed thatBos might have evolved in Africa and migrated to Eurasia during the Middle Pleistocene.[27] Middle Pleistocene aurochs fossils were also excavated in aSaharanerg in theHoggar Mountains.[28]
Fossils of the Indian subspecies (Bos primigenius namadicus) were excavated inalluvial deposits inSouth India dating to the Middle Pleistocene.[29] Remains of aurochs are common in Late Pleistocene sites across the Indian subcontinent.[30]
The earliest fossils in Europe date to the Middle Pleistocene. One site widely historically suggested to represent the first appearance of aurochs in Europe was the Notarchirico site in southern Italy, dating around 600,000 years ago,[31] however a 2024 re-examination of the site found that presence of aurochs at the locality was unsupported, with the oldest records of aurochs now placed at the Ponte Molle site in central Italy,[32] dating to around 550-450,000 years ago.[33] Aurochs were present in Britain byMarine Isotope Stage 11 ~400,000 years ago.[34]
The earliest remains aurochs in East Asia are uncertain, but may date to the late Middle Pleistocene.[35]
Late Pleistocene aurochs fossils were found inAffad 23 in Sudan dating to 50,000 years ago when the climate in this region was more humid than during theAfrican humid period.[36]
Following the most recentdeglaciation, the range of the aurochs expanded into Denmark and southern Sweden at the beginning of the Holocene, around 12-11,000 years ago.[37]
Illustration by Sigismund von Herberstein captioned:Urus sum, polonis Tur, germanis Aurox; ignari Bisontis nomen dederant; translated: "I am Urus,Tur in Polish,Aurox in German; the ignorant ones gave me the name Bison"
According to a 16th-century description bySigismund von Herberstein, the aurochs was pitch-black with a grey streak along the back; his wood carving made in 1556 was based on a culled aurochs, which he had received inMazovia.[38] In 1827,Charles Hamilton Smith published an image of an aurochs that was based on anoil painting that he had purchased from a merchant inAugsburg, which is thought to have been made in the early 16th century.[39] This painting is thought to have shown an aurochs,[5][40] although some authors suggested it may have shown a hybrid between an aurochs and domestic cattle, or a Polish steer.[41] Contemporary reconstructions of the aurochs are based onskeletons and the information derived from contemporaneous artistic depictions and historic descriptions of the animal.[5]
Remains of aurochs hair were not known until the early 1980s.[42] Depictions show that the North African aurochs may have had a light saddle marking on its back.[40] Calves were probably born with a chestnut colour, and young bulls changed to black with a whiteeel stripe running down the spine, while cows retained a reddish-brown colour. Both sexes had a light-coloured muzzle, but evidence for variation in coat colour does not exist. Egyptian grave paintings show cattle with a reddish-brown coat colour in both sexes, with a light saddle, but the horn shape of these suggest that they may depict domesticated cattle.[5]Many primitive cattle breeds, particularly those from Southern Europe, display similar coat colours to the aurochs, including the black colour in bulls with a light eel stripe, a pale mouth, and similar sexual dimorphism in colour.[5][40] A feature often attributed to the aurochs is blond forehead hairs. According to historical descriptions of the aurochs, it had long and curly forehead hair, but none mentions a certain colour. Although the colour is present in a variety of primitive cattle breeds, it is probably a discolouration that appeared after domestication.[5]
Drawing based on an aurochs bull skeleton fromLund and a cow skeleton fromCambridge, with characteristic features of the aurochs
Speculative profile of an Indian aurochs
The proportions and body shape of the aurochs were strikingly different from many modern cattle breeds. For example, the legs were considerably longer and more slender, resulting in a shoulder height that nearly equalled the trunk length. The skull, carrying the large horns, was substantially larger and more elongated than in most cattle breeds. As in other wild bovines, the body shape of the aurochs was athletic, and especially in bulls, showed a strongly expressed neck and shoulder musculature. Therefore, the fore hand was larger than the rear, similar to the wisent, but unlike many domesticated cattle. Even in carrying cows, the udder was small and hardly visible from the side; this feature is equal to that of other wild bovines.[5]
The aurochs was one of the largestherbivores inHolocene Europe. The size of an aurochs appears to have varied by region, with larger specimens in northern Europe than farther south. Aurochs in Denmark and Germany ranged in height at the shoulders between 155–180 cm (61–71 in) in bulls and 135–155 cm (53–61 in) in cows, while aurochs bulls inHungary reached 160 cm (63 in).[43]
The African aurochs was similar in size to the European aurochs in the Pleistocene, but declined in size during the transition to the Holocene; it may have also varied in size geographically.[44]
The body mass of aurochs appears to have shown some variability. Some individuals reached around 700 kg (1,540 lb), whereas those from the late Middle Pleistocene are estimated to have weighed up to 1,500 kg (3,310 lb).[5] The aurochs exhibited considerablesexual dimorphism in the size of males and females.[45]
The horns were massive, reaching 80 cm (31 in) in length and between 10 and 20 cm (3.9 and 7.9 in) in diameter.[40] Its horns grew from the skull at a 60-degree angle to the muzzle facing forwards and were curved in three directions, namely upwards and outwards at the base, then swinging forwards and inwards, then inwards and upwards. The curvature of bull horns was more strongly expressed than horns of cows.[5] The basal circumference ofhorn cores reached 44.5 cm (17.5 in) in the largest Chinese specimen and 48 cm (19 in) in a French specimen.[46] Some cattle breeds still show horn shapes similar to that of the aurochs, such as the Spanish fighting bull, and occasionally also individuals of derived breeds.[5]
A well-preserved aurochs bone yielded sufficient mitochondrial DNA for asequence analysis in 2010, which showed that its genome consists of 16,338base pairs.[47] Further studies using the aurochs whole genome sequence have identified candidate microRNA-regulateddomestication genes.[48] A comprehensive sequence analysis of Late Pleistocene and Holocene aurochs published in 2024 suggested that Indian aurochs (represented by modern zebu cattle) were the most genetically divergent aurochs population, having diverged from other aurochs around 300–166,000 years ago, with other aurochs populations spanning Europe and the Middle East to East Asia sharing much more recent common ancestry within the last 100,000 years. Late Pleistocene European aurochs were found to have a small (~3%) ancestry component from a divergent lineage that split prior to the divergence of Indian and other aurochs, suggested to be residual from earlier European aurochs populations. Towards the end of the Late Pleistocene, European aurochs experienced considerable gene flow from Middle Eastern aurochs. European Holocene aurochs primarily descend from those that were present in the Iberian Peninsula during theLast Glacial Maximum, with the Holocene also seeing mixing between previously isolated aurochs populations.[49]
Landscapes in Europe probably consisted of dense forests throughout much of the last few thousand years. The aurochs is likely to have usedriparian forests andwetlands along lakes.[45] Analysis of specimens found in Britain suggests that aurochs preferred inhabiting low lying relatively flat landscapes.[56]Pollen of mostly smallshrubs found in fossiliferoussediments with aurochs remains in China indicate that it preferredtemperate grassy plains orgrasslands borderingwoodlands.[46] It may have also lived in open grasslands.[57] In the warmAtlantic period of the Holocene, it was restricted to remaining open country and forest margins, where competition with livestock and humans gradually increased leading to a successive decline of the aurochs.[58]
Aurochs formed small herds mainly in winter, but typically lived singly or in smaller groups during the summer.[50] If aurochs had social behaviour similar to their descendants, social status would have been gained through displays and fights, in which both cows and bulls engaged.[40] Since it has ahypsodont jaw, it has been suggested to have been agrazer, with a food selection very similar to domesticated cattle[5] feeding on grass, twigs andacorns.[50]Mesowear analysis of Holocene Danish aurochspremolar teeth indicates that it changed from anabrasion-dominated grazer in the DanishPreboreal to a mixed feeder in theBoreal,Atlantic andSubboreal periods.[58]Dental microwear and mesowear analysis of specimens from the Pleistocene of Britain has found these aurochs had mixed feeding tobrowsing diets, rather than being strict grazers.[59]
Mating season was in September, and calves were born in spring.[50] Rutting bulls had violent fights, and evidence from theJaktorów forest shows that they were fully capable of mortally wounding one another. In autumn, aurochs fed for the winter, gaining weight and possessing a shinier coat than during the rest of the year. Calves stayed with their mothers until they were strong enough to join and keep up with the herd on the feeding grounds. Aurochs calves would have been vulnerable to predation bygrey wolves (Canis lupus) andbrown bears (Ursus arctos), while the immense size and strength of healthy adult aurochs meant they likely did not need to fear most predators.[5] According to historical descriptions, the aurochs was swift despite its build, could be very aggressive if provoked, and was not generally fearful of humans.[5] In Middle Pleistocene Europe, aurochs were likely predated upon by the "European jaguar"Panthera gombaszoegensis and thescimitar toothed-cat (Homotherium latidens),[60] with evidence for the consumption of aurochs bycave hyenas (Crocuta (Crocuta)spelaea) having been found from Late Pleistocene Italy.[61] Thelion (Panthera leo),tiger (Panthera tigris) and wolf are thought to have been the aurochs main predators during the Holocene.[45]
Acheulean layers inHunasagi on India's southernDeccan Plateau yielded aurochs bones with cut marks.[62] An aurochs bone with cut marks induced withflint was found in aMiddle Paleolithic layer at theNesher Ramla Homo site in Israel; it was dated toMarine Isotope Stage 5 about 120,000 years ago.[63] An archaeological excavation in Israel found traces of a feast held by theNatufian culture around 12,000 years BP, in which three aurochs were eaten. This appears to be an uncommon occurrence in the culture and was held in conjunction with the burial of an older woman, presumably of some social status.[64]Petroglyphs depicting aurochs inGobustan Rock Art in Azerbaijan date to theUpper Paleolithic toNeolithic periods.[65]Aurochs bones and skulls found at the settlements ofMureybet,Hallan Çemi andÇayönü indicate that people stored and shared food in thePre-Pottery Neolithic B culture.[66]Remains of an aurochs were also found in anecropolis inSidon, Lebanon, dating to around 3,700 years BP; the aurochs was buried together with numerous animals, a few human bones and foods.[67]
Petroglyphs depicting aurochs found inQurta in the upper Nile valley were dated to the Late Pleistocene about 19–15,000 years BP usingluminescence dating and are the oldest engravings found to date in Africa.[73] Aurochs are part of hunting scenes inreliefs in a tomb atThebes, Egypt dating to the 20th century BC, and in themortuary temple ofRamesses III atMedinet Habu dating to around 1175 BC. The latter is the youngest depiction of aurochs inAncient Egyptian art to date.[74]
Evidence has been found for the butchery of aurochs by archaic hominins in Europe during theMiddle Palaeolithic, such as the Biache-Saint-Vaast site in northern France dating to around 240,000 years ago, where bones of aurochs have been found burnt by fire and with cut marks, thought to have been created byNeanderthals.[75][76] At the late Middle Palaeolithic Cueva Des-Cubierta site in Spain, Neanderthals are proposed to have kept the skulls of aurochs as hunting trophies.[77]
The aurochs is widely represented inUpper Paleolithiccave paintings in theChauvet andLascaux caves in southern France dating to 36,000 and 21,000 years BP, respectively.[78]Two Paleolithicrock engravings in theCalabrianRomito Cave depict an aurochs.[79]Palaeolithic engravings showing aurochs were also found in theGrotta del Genovese on the Italian island ofLevanzo.[80]Upper Paleolithic rock engravings and paintings depicting the aurochs were also found in caves on theIberian Peninsula dating from theGravettian to theMagdalenian cultures.[81][82][83]Aurochs bones with chop and cut marks were found at variousMesolithic hunting and butchering sites in France, Luxemburg, Germany, the Netherlands, England and Denmark.[84] Aurochs bones were also found in Mesolithic settlements by theNarva andEmajõgi rivers inEstonia.[85] Aurochs and human bones were uncovered from pits andburnt mounds at several Neolithic sites in England.[86]A cup found in the Greek site ofVaphio shows a hunting scene, in which people try to capture an aurochs.[87] One of the bulls throws one hunter on the ground while attacking the second with its horns. The cup seems to date toMycenaean Greece.[88][89]Greeks andPaeonians hunted aurochs and used their huge horns as trophies, cups for wine, and offerings to the gods and heroes. The ox mentioned by Samus,Philippus of Thessalonica andAntipater as killed byPhilip V of Macedon on the foothills of mountainOrvilos, was actually an aurochs; Philip offered the horns, which were 105 cm (41 in) long and the skin to a temple ofHercules.[90]The aurochs was described inJulius Caesar'sCommentarii de Bello Gallico.[6]Aurochs were occasionally captured and exhibited invenatio shows inRoman amphitheatres such as theColosseum.[91] Aurochs horns were often used by Romans as hunting horns.[5]
Aurochs were hunted with arrows, nets and hunting dogs, and its hair on the forehead was cut from the living animal; belts were made out of this hair and believed to increase thefertility of women. When the aurochs was slaughtered, theos cordis was extracted from the heart; this bone contributed to the mystique and magical powers that were attributed to it.[5]In eastern Europe, the aurochs has left traces in expressions like "behaving like an aurochs" for a drunken person behaving badly, and "a bloke like an aurochs" for big and strong people.[45]
The earliest-knowndomestication of the aurochs dates to theNeolithic Revolution in theFertile Crescent, where cattle hunted and kept by Neolithic farmers gradually decreased in size between 9800 and 7500 BC. Aurochs bones found atMureybet andGöbekli Tepe are larger in size than cattle bones from later Neolithic settlements in northernSyria likeDja'de el-Mughara andTell Halula.[96]InLate Neolithic sites of northernIraq and westernIran dating to the sixth millennium BC, cattle remains are also smaller but more frequent, indicating that domesticated cattle were imported during theHalaf culture from the central Fertile Crescent region.[97]Results of genetic research indicate that the moderntaurine cattle (Bos taurus) arose from 80 aurochstamed in southeasternAnatolia and northern Syria about 10,500 years ago.[15]Taurine cattle spread into theBalkans and northern Italy along theDanube River and the coast of theMediterranean Sea.[98]Hybridisation between male aurochs and early domestic cattle occurred in central Europe between 9500 and 1000 BC.[99]Analyses of mitochondrial DNA sequences of Italian aurochs specimens dated to 17–7,000 years ago and 51 modern cattle breeds revealed some degree ofintrogression of aurochs genes into south European cattle, indicating that female aurochs had contact with free-ranging domestic cattle.[100] Cattle bones of various sizes found at aChalcolithic settlement in theKutná Hora District provide further evidence for hybridisation of aurochs and domestic cattle between 3000 and 2800 BC in theBohemian region.[43]Whole genome sequencing of a 6,750-year-old aurochs bone found in England was compared with genome sequence data of 81 cattle andsingle-nucleotide polymorphism data of 1,225 cattle. Results revealed that British and Irish cattle breeds share somegenetic variants with the aurochs specimen; early herders in Britain might have been responsible for the localgene flow from aurochs into the ancestors of British and Irish cattle.[101] TheMurboden cattle breed also exhibits sporadic introgression of female European aurochs into domestic cattle in theAlps.[102] Domestic cattle continued to diminish in both body and horn size until the Middle Ages.[87]
Comparative analysis ofsingle-nucleotide polymorphisms and sharedalleles revealed admixture between East Asian aurochs and introduced taurine cattle inancient China, for example atShimao. This suggested the incorporation of local aurochs into domestic cattle as far back as 4,000 years BP, either through spontaneousintrogression, or the capture of different aurochs groups to supplement domestic stocks. The same study detected derived alleles shared by aurochs and modern taurine cattle in East Asia, especially amongTibetan breeds. Introgression with local aurochs could have facilitated rapid adaptation to new environments.[17]
The Indian aurochs is thought to have been domesticated 10,000–8,000 years ago.[103]Aurochs fossils found at the Neolithic site ofMehrgarh in Pakistan are dated to around 8,000 years BP and represent some of the earliest evidence for its domestication on the Indian subcontinent.[30] Female Indian aurochs contributed to the gene pool ofzebu (Bos indicus) between 5,500 and 4,000 years BP during the expansion ofpastoralism in northern India. The zebu initially spread eastwards toSoutheast Asia.[104]Hybridisation between zebu and early taurine cattle occurred in theNear East after 4,000 years BP coinciding with thedrought period during the4.2-kiloyear event.[105] The zebu was introduced toEast Africa about 3,500–2,500 years ago,[98] and reachedMongolia in the 13th and 14th centuries.[106]
A third domestication event thought to have occurred inEgypt's Western Desert is not supported by results of an analysis ofgenetic admixture, introgression and migration patterns of 3,196 domestic cattle representing 180 populations. However, the same study supported extensive hybridization between taurine cattle in Africa, arrived from the Near East after domestication, and local wild African aurochs prior to the entry of the zebu in Africa.[98] The zebu was introduced throughAncient Egypt and started to spread comprehensively throughWest Africa in the last 1,400 years, along withArabic cultural influences. Most modernAfrican cattle breeds are hybridized to a variable extent with Indicine cattle, with introgression being most reduced in areas of West Africa where thetse-tse fly is present.[107]
Scene from the tombTT60 inThebes, Egypt, depicting a North African aurochs bull, cow and calf being hunted withdogs andjavelins.
The Indian aurochs (B. p. namadicus) became extinct sometime during the Holocene period, likely due to habitat loss caused by expandingpastoralism and interbreeding with the domesticzebu.[104][108] The timing of extinction of aurochs in the Indian subcontinent is unclear, due to difficulty distinguishing aurochs remains from those of domestic cattle, with a 2021 review suggesting remains from Mehrgarh, Pakistan, dating to around 8,000 years ago "might constitute the only dated and reliably identified evidence" of Holocene Indian aurochs.[30] The extinction probably predates the historical period, due to a lack of references to the aurochs in Indian texts.[109]
A 2014 review suggested that the youngest remains of African aurochs (B. p. mauritanicus) dated to around 6,000 yearsBefore Present (BP),[110] though some authors suggest that it may have survived until at least to theRoman period, as indicated by remains found inButo andFaiyum in theNile Delta.[44]
In China, aurochs persisted until at least 3,600 BP.[111]
The Eurasian aurochs (B. p. primigenius) was present in southern Sweden during theHolocene climatic optimum until at least 7,800 years BP.[112] In Denmark, the first-knownlocal extinction of the aurochs occurred after thesea level rise on the newly formedDanish islands about 8,000–7,500 years BP, and the last documented aurochs lived in southernJutland around 3,000 years BP.[37] The latest-known aurochs fossil inGreat Britain dates to 3,245 years BP, and it was probably extinct by 3,000 years ago.[113]
Excessive hunting began and continued until the aurochs was nearly extinct. The gradual extinction of the aurochs inCentral Europe was concurrent with theclearcutting of large forest tracts between the 9th and 12th centuries.[50]
By the 13th century, the aurochs existed only in small numbers inEastern Europe, and hunting it became a privilege of nobles and later royals.[5] The population inHungary was declining from at least the 9th century and was extinct in the 13th century.[114][115]
Findings fromsubfossil records indicate that wild aurochs might have survived in northwesternTransylvania until the 14th to 16th century, inwestern Moldavia until probably the early 17th century.[116][117]
The last-known aurochs herd lived in amarshy woodland in Poland'sJaktorów Forest. It decreased from around 50 individuals in the mid 16th century to four individuals by 1601. The last aurochs cow died in 1627 from natural causes.[118]
A 2021 study argued that the aurochs possibly survived in northeasternBulgaria until at least the 17th century.[119] A horn-core excavated in 2020 inSofia was identified as being from an aurochs; the archaeological layer in which it was found was dated to the second half of the 17th or first half of the 18th century, suggesting that aurochs may have survived in Bulgaria until that date.[120]
In the early 1920s,Heinz Heck initiated aselective breeding program inHellabrunn Zoo attempting tobreed back the aurochs using several cattle breeds; the result is calledHeck cattle.[121]Herds of these cattle were released toOostvaardersplassen, apolder in the Netherlands in the 1980s as aurochs surrogates for naturalistic grazing with the aim to restore prehistorical landscapes.[122] Large numbers of them died ofstarvation during the cold winters of 2005 and 2010, and the project of no interference ended in 2018.[123]
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