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=== Evolution === [[Molecular clock#Calibration|Calibrations]] using [[fossil]]s of 16 Bovidae species indicate that the [[Bovini]] tribe evolved about {{mya|11.7}}.<ref>{{cite journal |last=Bibi |first=F. |year=2013 |title=A multi-calibrated mitochondrial phylogeny of extant Bovidae (Artiodactyla, Ruminantia) and the importance of the fossil record to systematics |journal=BMC Evolutionary Biology |volume=13 |issue=1 |pages=166 |bibcode=2013BMCEE..13..166B |doi=10.1186/1471-2148-13-166 |pmc=3751017 |pmid=23927069 |doi-access=free}}</ref> The ''[[Bos]]'' and ''[[Bison]]'' [[genetic lineage]]s are estimated to have [[Genetic divergence|genetically diverged]] from the Bovini about {{mya|2.5|1.65}}.<ref name=Wang2018>{{cite journal |author1=Wang, K. |name-list-style=amp |author2=Lenstra, J.A. |author3=Liu, L. |author4=Hu, Q. |author5=Ma, T. |author6=Qiu, Q. |author7=Liu, J. |year=2018 |title=Incomplete lineage sorting rather than hybridization explains the inconsistent phylogeny of the wisent |journal=Communications Biology |volume=1 |issue=1 |page=169 |doi=10.1038/s42003-018-0176-6 |pmid=30374461 |pmc=6195592 |doi-access=free}}</ref><ref>{{cite journal |author1=Zeyland, J. |name-list-style=amp |author2=Wolko, Ł. |author3=Lipiński, D. |author4=Woźniak, A. |author5=Nowak, A. |author6=Szalata, M. |author7=Bocianowski, J. |author8=Słomski, R. |year=2012 |title=Tracking of wisent–bison–yak mitochondrial evolution |journal=Journal of Applied Genetics |volume=53 |issue=3 |pages=317–322 |doi=10.1007/s13353-012-0090-4 |pmid=22415349 |pmc=3402669 |doi-access=free}}</ref> The following [[cladogram]] shows the [[Phylogenetics|phylogenetic]] relationships of the aurochs based on analysis of [[Nuclear DNA|nuclear]] and [[mitochondrial DNA|mitochondrial]] genomes in the Bovini [[Tribe (biology)|tribe]]:<ref name=Wang2018/><ref>{{cite journal |last=Bibi |first=F. |title=A multi-calibrated mitochondrial phylogeny of extant Bovidae (Artiodactyla, Ruminantia) and the importance of the fossil record to systematics |journal=BMC Evolutionary Biology |date=2013 |volume=13 |issue=1 |pages=166 |doi=10.1186/1471-2148-13-166 |pmid=23927069 |pmc=3751017 |doi-access=free |bibcode=2013BMCEE..13..166B}}</ref><ref>{{Cite journal |last1=Sinding |first1=M.-H. S. |last2=Ciucani |first2=M. M. |last3=Ramos-Madrigal |first3=J. |last4=Carmagnini |first4=A. |last5=Rasmussen |first5=J. A. |last6=Feng |first6=S. |last7=Chen |first7=G. |last8=Vieira |first8=F. G. |last9=Mattiangeli |first9=V. |last10=Ganjoo |first10=R. K. |last11=Larson |first11=G. |last12=Sicheritz-Pontén |first12=T. |last13=Petersen |first13=B. |last14=Frantz |first14=L. |last15=Gilbert |first15=M. T. P. |date=2021 |title=Kouprey (''Bos sauveli'') genomes unveil polytomic origin of wild Asian Bos |journal=iScience |volume=24 |issue=11 |pages=103226 |bibcode=2021iSci...24j3226S |doi=10.1016/j.isci.2021.103226 |pmc=8531564 |pmid=34712923}}</ref> {{clade |label1=[[Bovini]] |1={{clade |1=[[Bubalina]] (buffalo) |label2=''[[Bos]]'' |2={{clade |1= '''''Bos primigenius''''' (aurochs) |2={{clade |1={{clade |1=''[[Bos mutus]]'' (wild yak) |2={{clade |1=''[[Bison bison]]'' (American bison) |2=''[[Bison bonasus]]'' (European bison/wisent) }}}} |2={{clade |1=''[[Bos javanicus]]'' (banteng) |2=''[[Bos gaurus]]'' (gaur) |3=''[[Bos sauveli]]'' (kouprey) }}}}}}}}}} The cold [[Pliocene climate]] caused an extension of open [[grassland]], which enabled the [[evolution]] of large grazers.<ref name="vVuure05">{{cite book |title=Retracing the Aurochs: History, Morphology and Ecology of an extinct wild Ox |last=Van Vuure, C. |year=2005 |location=Sofia |publisher=Pensoft Publishers |isbn=954-642-235-5}}</ref> 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 was [[Excavation (archaeology)|excavated]] in the [[Sivalik Hills]] in India that dates to the [[Early Pleistocene]] about {{mya|2}}.<ref>{{cite journal |author1=Samartzidou, E. |name-list-style=amp |author2=Pandolfi, L. |year=2021 |title=''Bos primigenius'' Bojanus, 1827 (Mammalia, Bovidae) in Greece: new finds and a revision of the species, with a comparison with body-size variations of aurochs from the Italian Peninsula |journal=Acta Zoologica Bulgarica |volume=74 |pages=119–139 |author3=Tsoukala, E. |author4=Maniatis, Y. |author5=Stoulos, S.}}</ref> An aurochs skull excavated in Tunisia's [[Kef Governorate]] from early [[Middle Pleistocene]] [[strata]] dating about {{mya|0.78}} is the oldest well-dated fossil specimen to date. The authors of the study proposed that ''Bos'' might have evolved in Africa and migrated to Eurasia during the Middle Pleistocene.<ref>{{cite journal |author1=Martínez-Navarro, B.|name-list-style=amp |author2=Karoui-Yaakoub, N. |author3=Oms, O. |author4=Amri, L. |author5=López-García, J.M. |author6=Zerai, K. |author7=Blain, H.A. |author8=Mtimet, M.S. |author9=Espigares, M.P. |author10=Ali, N.B.H. |author11=Ros-Montoya, S. |author12=Boughdiri, M. |author13=Agustí J. |author14=Khayati-Ammar, H. |author15=Maalaoui K. |author16=El Khir, M.O. |author17=Sala, R. |author18=Othmani, A. |author19=Hawas, R. |author20=Gómez-Merino, G. |author21=Solè, À. |author22=Carbonell, E. |author23=Palmqvist, P. |year=2014 |title=The early Middle Pleistocene archeopaleontological site of Wadi Sarrat (Tunisia) and the earliest record of ''Bos primigenius'' |journal=Quaternary Science Reviews |volume=90 |pages=37–46 |doi=10.1016/j.quascirev.2014.02.016|bibcode=2014QSRv...90...37M}}</ref> Middle Pleistocene aurochs fossils were also excavated in a [[Sahara]]n [[Erg (landform)|erg]] in the [[Hoggar Mountains]].<ref>{{cite book |last=Thomas |first=H. |title=Géologie et paléontologie du gisement acheuléen de l'erg Tihodaïne, Ahaggar Sahara Algérien |publisher=Memoires du centre de recherches anthlropologiques, prehistoriques et ethnographiques |year=1977 |location=Paris}}</ref> Fossils of the Indian subspecies (''Bos primigenius namadicus'') were excavated in [[alluvial deposit]]s in [[South India]] dating to the Middle Pleistocene.<ref>{{cite journal |last=Pilgrim |first=G.E. |year=1947 |title=The evolution of the buffaloes, oxen, sheep and goats |journal=Zoological Journal of the Linnean Society |volume=41 |issue=279 |pages=272–286 |doi=10.1111/j.1096-3642.1940.tb02077.x}}</ref> Remains of aurochs are common in Late Pleistocene sites across the Indian subcontinent.<ref name="Turvey_al2021" /> 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,<ref>{{Cite journal |last1=Gómez-Olivencia |first1=Asier |last2=Sala |first2=Nohemi |last3=Arceredillo |first3=D. |last4=García |first4=N. |last5=Martínez-Pillado |first5=V. |last6=Rios-Garaizar |first6=J. |last7=Garate |first7=D. |last8=Solar |first8=G. |last9=Libano |first9=I. |date=2015 |title=The Punta Lucero Quarry site (Zierbena, Bizkaia): a window into the Middle Pleistocene in the Northern Iberian Peninsula |journal=Quaternary Science Reviews |volume=121 |pages=52–74 |doi=10.1016/j.quascirev.2015.05.001|bibcode=2015QSRv..121...52G}}</ref> 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,<ref>{{Cite journal |last1=Mecozzi |first1=B. |last2=Iannucci |first2=A. |last3=Carpentieri |first3=M. |last4=Pineda |first4=A. |last5=Rabinovich |first5=R. |last6=Sardella |first6=R. |last7=Moncel |first7=M.-H. |date=2024 |title=Climatic and environmental changes of ~100 thousand years: The mammals from the early Middle Pleistocene sequence of Notarchirico (southern Italy) |journal=PLOS ONE |volume=19 |issue=10 |pages=e0311623 |doi=10.1371/journal.pone.0311623 |pmc=11498728 |pmid=39441829 |doi-access=free|bibcode=2024PLoSO..1911623M }}</ref> dating to around 550–450,000 years ago.<ref>{{Cite journal |last1=Mecozzi |first1=B. |last2=Iannucci |first2=A. |last3=Mancini |first3=M. |last4=Sardella |first4=R. |date=2021 |title=Redefining Ponte Molle (Rome, central Italy): an important locality for Middle Pleistocene mammal assemblages of Europe |journal=Alpine and Mediterranean Quaternary |volume=34 |issue=1 |pages=131–154 |doi=10.26382/AMQ.2021.09}}</ref> Aurochs were present in Britain by [[Marine Isotope Stage 11]] ~400,000 years ago.<ref>{{Cite journal |last1=Preece |first1=R.C. |last2=Parfitt |first2=S.A. |last3=Bridgland |first3=D.R. |last4=Lewis |first4=S.G. |last5=Rowe |first5=P.J. |last6=Atkinson |first6=T.C. |last7=Candy |first7=I. |last8=Debenham |first8=N.C. |last9=Penkman |first9=K.E.H. |last10=Rhodes |first10=E.J. |last11=Schwenninger |first11=J.-L. |last12=Griffiths |first12=H.I. |last13=Whittaker |first13=J.E. |last14=Gleed-Owen |first14=C. |date=2007 |title=Terrestrial environments during MIS 11: evidence from the Palaeolithic site at West Stow, Suffolk, UK |journal=Quaternary Science Reviews |volume=26 |issue=9–10 |pages=1236–1300 |doi=10.1016/j.quascirev.2006.11.016 |bibcode=2007QSRv...26.1236P}}</ref> The earliest remains aurochs in East Asia are uncertain, but may date to the late Middle Pleistocene.<ref>{{cite journal |author1=Tong, H. |name-list-style=amp |author2=Chen, X. |year=2018 |title=New fossils of ''Bos primigenius'' (Artiodactyla, Mammalia) from Nihewan and Longhua of Hebei, China |journal=Vertebrata PalAsiatica |volume=56 |issue=1 |pages=69–92 |author3=Zhang, B. |author4=Wang, F.}}</ref> Late Pleistocene aurochs fossils were found in [[Affad 23]] in Sudan dating to 50,000 years ago when the climate in this region was more humid than during the [[African humid period]].<ref>{{cite journal |author1=Osypinska, M. |name-list-style=amp |author2=Osypinski, P. |author3=Belka, Z. |author4=Chlodnicki, M. |author5=Wiktorowicz, P. |author6=Ryndziewicz, R. |author7=Kubiak, M. |year=2021 |title=Wild and Domestic Cattle in the Ancient Nile Valley: Marks of ecological change |journal=Journal of Field Archaeology |volume=46 |issue=7 |pages=429–447 |doi=10.1080/00934690.2021.1924491 |s2cid=236373843}}</ref> Following the most recent [[deglaciation]], the range of the aurochs expanded into Denmark and southern Sweden at the beginning of the Holocene, around 12–11,000 years ago.<ref name="Gravlund_al2012">{{cite journal |author1=Gravlund, P. |author2=Aaris-Sørensen, K. |author3=Hofreiter, M. |author4=Meyer, M. |author5=Bollback, J.P. |author6=Noe-Nygaard, N |name-list-style=amp |year=2012 |title=Ancient DNA extracted from Danish aurochs (''Bos primigenius''): genetic diversity and preservation |journal=Annals of Anatomy |volume=194 |issue=1 |pages=103–111 |citeseerx=10.1.1.392.4989 |doi=10.1016/j.aanat.2011.10.011 |pmid=22188739}}</ref>
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