Editing Bison (section)

==Evolution and genetic history==
The bovine tribe (Bovini) split about 5 to 10 million years ago into the buffalos (''[[Bubalus]]'' and ''[[Syncerus]]'') and a group leading to bison and taurine cattle.<ref>{{cite journal |title=Maternal and Paternal Lineages in Cross-Breeding Bovine Species. Has Wisent a Hybrid Origin? |journal=Molecular Biology and Evolution |volume=21 |issue=7 |pages=1165–1170 |date=22 January 2004 |doi=10.1093/molbev/msh064 |pmid=14739241 |last1=Verkaar |first1=Edward L. C. |last2=Nijman |first2=Isaäc J. |last3=Beeke |first3=Maurice |last4=Hanekamp |first4=Eline |last5=Lenstra |first5=Johannes A.|doi-access=free }}</ref> Genetic evidence from [[nuclear DNA]] indicates that the closest living relatives of bison are [[Wild yak|yaks]], with bison being nested within the genus ''[[Bos]],'' rendering ''Bos'' without including bison [[paraphyletic]]. While nuclear DNA indicates that both extant bison species are each other's closest living relatives, the [[mitochondrial DNA]] of European bison is more closely related to that of domestic cattle and [[aurochs]] (while the mitochondrial DNA of American bison is closely related to that of yaks). This discrepancy is either suggested to be the result of [[incomplete lineage sorting]] or ancient [[introgression]].<ref name="WangEtal2018">Wang, K., Lenstra, J. A., Liu, L., Hu, Q., Ma, T., Qiu, Q., & Liu, J. (2018). Incomplete lineage sorting rather than hybridization explains the inconsistent phylogeny of the wisent. Communications biology, 1(1), 1–9.</ref><ref name=":4">{{Cite journal|last1=Grange|first1=Thierry|last2=Brugal|first2=Jean-Philip|last3=Flori|first3=Laurence|last4=Gautier|first4=Mathieu|last5=Uzunidis|first5=Antigone|last6=Geigl|first6=Eva-Maria|date=September 2018|title=The Evolution and Population Diversity of Bison in Pleistocene and Holocene Eurasia: Sex Matters|journal=Diversity|language=en|volume=10|issue=3|pages=65|doi=10.3390/d10030065|doi-access=free}}</ref><ref>{{Cite journal |last1=Zver |first1=Lars |last2=Toškan |first2=Borut |last3=Bužan |first3=Elena |date=September 2021 |title=Phylogeny of Late Pleistocene and Holocene Bison species in Europe and North America |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618221002421 |journal=Quaternary International |language=en |volume=595 |pages=30–38 |doi=10.1016/j.quaint.2021.04.022|bibcode=2021QuInt.595...30Z }}</ref> Bison are widely believed to have evolved from a lineage belonging to the extinct genus ''[[Leptobos]]'' during the [[Piacenzian|Late Pliocene]] to [[Early Pleistocene]] in Asia. The earliest members of the bison lineage, known from the Late Pliocene to Early Pleistocene of the Indian Subcontinent (''Bison sivalensis'') and China (''Bison palaeosinensis''), approximately 3.4-2.6 million years ago (Ma) are placed in the subgenus ''Bison'' (''Eobison'')''.'' The oldest remains of ''Eobison'' in Europe are those ''Bison georgicus'' found in [[Dmanisi historic site|Dmanisi]], Georgia, dated to around 1.76 Ma. More [[Primitive (phylogenetics)|derived]] members of the genus are placed in the subgenus ''Bison'' (''Bison''), which first appeared towards the end of the Early Pleistocene, around 1.2 Ma, with early members of the subgenus including the widespread ''[[Bison schoetensacki]]''.<ref name=":2">{{Cite journal|last1=Sorbelli|first1=Leonardo|last2=Alba|first2=David M.|last3=Cherin|first3=Marco|last4=Moullé|first4=Pierre-Élie|last5=Brugal|first5=Jean-Philip|last6=Madurell-Malapeira|first6=Joan|date=2021-06-01|title=A review on Bison schoetensacki and its closest relatives through the early-Middle Pleistocene transition: Insights from the Vallparadís Section (NE Iberian Peninsula) and other European localities|url=https://www.sciencedirect.com/science/article/pii/S0277379121001402|journal=Quaternary Science Reviews|language=en|volume=261|pages=106933|doi=10.1016/j.quascirev.2021.106933|bibcode=2021QSRv..26106933S |s2cid=235527116 |issn=0277-3791}}</ref>

The [[steppe bison]] (''Bison priscus'') first appeared during the mid-[[Middle Pleistocene]] in eastern Eurasia, and subsequently became widely distributed across Eurasia.<ref name=":2" /> During the late Middle Pleistocene, around 195,000-135,000 years ago, the steppe bison migrated across the [[Bering land bridge]] into North America, becoming ancestral to North American bison species, including the large ''[[Bison latifrons]],'' and the smaller ''[[Bison antiquus]],'' which became extinct at the end of the Late Pleistocene.<ref>{{Cite journal|last1=Froese|first1=Duane|last2=Stiller|first2=Mathias|last3=Heintzman|first3=Peter D.|last4=Reyes|first4=Alberto V.|last5=Zazula|first5=Grant D.|last6=Soares|first6=André E. R.|last7=Meyer|first7=Matthias|last8=Hall|first8=Elizabeth|last9=Jensen|first9=Britta J. L.|last10=Arnold|first10=Lee J.|last11=MacPhee|first11=Ross D. E.|date=2017-03-28|title=Fossil and genomic evidence constrains the timing of bison arrival in North America|journal=Proceedings of the National Academy of Sciences|language=en|volume=114|issue=13|pages=3457–3462|doi=10.1073/pnas.1620754114|issn=0027-8424|pmid=28289222|pmc=5380047 |bibcode=2017PNAS..114.3457F |doi-access=free }}</ref> Modern American bison are thought to have evolved from ''B.&nbsp;antiquus'' during the Late Pleistocene-Holocene transition via the intermediate form ''[[Bison occidentalis]]''.<ref name="Wilsonetal.2008">{{cite journal|last1=Wilson|first1=M.C.|last2=Hills|first2=L.V.|last3=Shapiro|first3=B.|year=2008|title=Late Pleistocene northward-dispersing ''Bison antiquus'' from the Bighill Creek Formation, Gallelli Gravel Pit, Alberta, Canada, and the fate of ''Bison occidentalis''|journal=Canadian Journal of Earth Sciences|volume=45|issue=7|pages=827–59|bibcode=2008CaJES..45..827W|doi=10.1139/E08-027}}</ref> The European bison, ''Bison bonasus,'' first appeared in Europe during the late Middle Pleistocene, where it existed in [[sympatry]] with the steppe bison. Its relationship with other extinct bison species is unclear, though it appears to be only distantly related to the steppe and American bisons, with possibly some interbreeding between the two lineages during the Middle Pleistocene.<ref name=":4" /> The steppe bison survived into the early-mid [[Holocene]] in Alaska-Yukon and eastern Siberia, before becoming extinct.<ref>{{Cite journal|last1=Boeskorov|first1=Gennady G.|last2=Potapova|first2=Olga R.|last3=Protopopov|first3=Albert V.|last4=Plotnikov|first4=Valery V.|last5=Agenbroad|first5=Larry D.|last6=Kirikov|first6=Konstantin S.|last7=Pavlov|first7=Innokenty S.|last8=Shchelchkova|first8=Marina V.|last9=Belolyubskii|first9=Innocenty N.|last10=Tomshin|first10=Mikhail D.|last11=Kowalczyk|first11=Rafal|date=June 2016|title=The Yukagir Bison: The exterior morphology of a complete frozen mummy of the extinct steppe bison, Bison priscus from the early Holocene of northern Yakutia, Russia|url=https://linkinghub.elsevier.com/retrieve/pii/S1040618215012100|journal=Quaternary International|language=en|volume=406|pages=94–110|doi=10.1016/j.quaint.2015.11.084|bibcode=2016QuInt.406...94B |s2cid=133244037 }}</ref><ref>{{Cite journal|last1=Zazula|first1=Grant D.|last2=Hall|first2=Elizabeth|last3=Hare|first3=P. Gregory|last4=Thomas|first4=Christian|last5=Mathewes|first5=Rolf|last6=La Farge|first6=Catherine|last7=Martel|first7=André L.|last8=Heintzman|first8=Peter D.|last9=Shapiro|first9=Beth|date=November 2017|title=A middle Holocene steppe bison and paleoenvironments from the Versleuce Meadows, Whitehorse, Yukon, Canada|url=http://www.nrcresearchpress.com/doi/10.1139/cjes-2017-0100|journal=Canadian Journal of Earth Sciences|language=en|volume=54|issue=11|pages=1138–1152|doi=10.1139/cjes-2017-0100|bibcode=2017CaJES..54.1138Z |hdl=1807/78639 |s2cid=54951935 |issn=0008-4077|hdl-access=free}}</ref>

[[File:Euroameribison.jpg|thumb|Skulls of European bison (left) and American bison (right)]]

Prior to the late 19th century, the population of American bison likely numbered in the tens of millions, perhaps as many as 60 million.<ref>{{Cite journal |last=Hedrick |first=P. W. |date=2009-07-01 |title=Conservation Genetics and North American Bison (Bison bison) |url=https://academic.oup.com/jhered/article-lookup/doi/10.1093/jhered/esp024 |journal=Journal of Heredity |language=en |volume=100 |issue=4 |pages=411–420 |doi=10.1093/jhered/esp024 |issn=0022-1503}}</ref> During the population bottleneck caused by the great slaughter of American bison during the 19th century, the number of bison remaining alive in North America declined to as low as 541. During that period, a handful of ranchers gathered remnants of the existing herds to save the species from extinction. These ranchers bred some of the bison with cattle in an effort to produce "cattleo"<ref name="Halbert2007" /> (today called "[[beefalo]]"). Accidental crossings were also known to occur. Generally, male domestic bulls were crossed with bison cows, producing offspring of which only the females were fertile. The crossbred animals did not demonstrate any form of [[Heterosis|hybrid vigor]], so the practice was abandoned. Wisent-American bison hybrids were briefly experimented with in Germany (and found to be fully fertile) and a herd of such animals is maintained in Russia. A herd of cattle-wisent crossbreeds ([[zubron]]) is maintained in Poland. First-generation crosses do not occur naturally, requiring caesarean delivery. First-generation males are infertile. The U.S. National Bison Association has adopted a code of ethics that prohibits its members from deliberately crossbreeding bison with any other species. In the United States, many ranchers are now using DNA testing to cull the residual cattle genetics from their bison herds. The proportion of cattle DNA that has been measured in introgressed individuals and bison herds today is typically quite low, ranging from 0.56 to 1.8%.<ref name=Halbert2007/><ref name=Polziehn/>

There are also remnant purebred American bison herds on public lands in North America. Two subspecies of bison exist in North America: the ''plains bison'' and the ''wood bison''.<ref>{{Cite web |title=Bison |url=https://www.thecanadianencyclopedia.ca/en/article/bison |access-date=2023-07-23 |website=www.thecanadianencyclopedia.ca |language=en}}</ref> Herds of importance are found in [[Yellowstone National Park]], [[Wind Cave National Park]] in [[South Dakota]], [[Blue Mounds State Park]] in [[Minnesota]], [[Elk Island National Park]] in [[Alberta]], and [[Grasslands National Park]] in [[Saskatchewan]]. In 2015, a purebred herd of 350 individuals was identified on public lands in the [[Henry Mountains]] of southern [[Utah]] via [[genetic testing]] of mitochondrial and nuclear DNA.<ref>{{cite journal |title=Genetic Analysis of the Henry Mountains Bison Herd |vauthors=Ranglack DH, Dobson LK, du Toit JT, Derr J|journal=PLOS ONE|date=17 December 2015 |doi=10.1371/journal.pone.0144239 |volume=10 |issue=12 |page=e0144239 |pmid=26673758 |pmc=4682953 |bibcode=2015PLoSO..1044239R|doi-access=free}}</ref> This study, published in 2015, also showed the [[Henry Mountains bison herd]] to be free of [[brucellosis]], a bacterial disease that was imported with non-native domestic cattle to North America.<ref>{{cite journal |title=On the Origin of Brucellosis in Bison of Yellowstone National Park: A Review |author=Mary Meagher, Margaret E. Meyer |journal=Conservation Biology |date=September 1994 |jstor=2386505 |volume=8 |issue=3 |pages=645–653 |doi=10.1046/j.1523-1739.1994.08030645.x|bibcode=1994ConBi...8..645M }}</ref>

In 2021, the [[American Society of Mammalogists]] considered ''Bison'' to be a subgenus, and placed both bison species back into ''[[Bos]]''.<ref>Groves, C. P. & Grubb, P. 2011. Ungulate taxonomy. The Johns Hopkins University Press, Baltimore, Maryland.|Wang, K., Lenstra, J. A., Liu, L., Hu, Q., Ma, T., Qiu, Q., & Liu, J. (2018). Incomplete lineage sorting rather than hybridization explains the inconsistent phylogeny of the wisent. Communications biology, 1(1), 1–9. Accessed at https://www.mammaldiversity.org/explore.html#species-id=1006256 {{Webarchive|url=https://web.archive.org/web/20201028140029/https://www.mammaldiversity.org/explore.html#species-id=1006256 |date=28 October 2020 }}.</ref>{{failed verification|reason=This source does not declare Bison to be a subgenus so much as moving the species directly into Bos|date=November 2022}}

Relationships of bovines based on nuclear DNA, after Sinding, et al. 2021.<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
|1=[[Bubalina]] (true buffalo)
|label2=''[[Bos]]''
|2={{clade
|1= ''[[Bos primigenius]]'' +''[[Bos taurus]]'' (aurochs and cattle)
|2={{clade
|1={{clade
|1=''[[Bos mutus]]'' (wild yak)
|label2=''Bison''
|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)
}}}}}}}}