Bovina is a subtribe of the

Bovini The tribe Bovini, or wild cattle, are medium to massive bovines that are native to North America, Eurasia, and Africa. These include the enigmatic, antelope-like saola, the African and Asiatic buffalos, and a clade that consists of bison and t ...

tribe that generally includes the two living genera, '' Bison'' and ''

Bos ''Bos'' (from Latin '' bōs'': cow, ox, bull) is the genus of wild and domestic cattle. ''Bos'' is often divided into four subgenera: ''Bos'', ''Bibos'', ''Novibos'', and ''Poephagus'', but including these last three divisions within the gen ...

''. However, this dichotomy has been challenged recently by molecular work that suggests that ''Bison'' should be regarded as a subgenus of ''Bos''. Wild bovinans can be found naturally in North America and Eurasia (although domestic and

feral A feral () animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in some ...

populations have been introduced worldwide).

Taxonomy

Placement within Bovini

The majority of phylogenetic work based on

ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chromo ...

chromosomal A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...

analysis, autosomal introns and mitochondrial DNA has recovered three distinctive subtribes of

: Pseudorygina (represent solely by the

saola The saola (''Pseudoryx nghetinhensis''), also called spindlehorn, Asian unicorn, or infrequently, Vu Quang bovid, is one of the world's rarest large mammals, a forest-dwelling bovine native to the Annamite Range in Vietnam and Laos. It was descr ...

Bubalina Bubalina is a subtribe of wild cattle that includes the various species of true buffalo. Species include the African buffalo, the anoas, and the wild water buffalo (including the domesticated variant water buffalo). Buffaloes can be found ...

(buffalo), and Bovina.

Genera and species

Extant

In 1945 American paleontologist and mammalogist

George Gaylord Simpson George Gaylord Simpson (June 16, 1902 – October 6, 1984) was an American paleontologist. Simpson was perhaps the most influential paleontologist of the twentieth century, and a major participant in the modern synthesis, contributing '' Tempo ...

had considered there to be three genera of bovinans. In addition to recognizing ''Bos'' and ''Bison'', he assigned the several Asiatic tropical species such as

gaur The gaur (''Bos gaurus''; ), also known as the Indian bison, is a bovine native to South Asia and Southeast Asia, and has been listed as Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 m ...

and banteng into the genus ''Bibos''. The

German German(s) may refer to: * Germany (of or related to) ** Germania (historical use) * Germans, citizens of Germany, people of German ancestry, or native speakers of the German language ** For citizens of Germany, see also German nationality law **Ge ...

zoologist Herwart Bohlken also agreed with these conclusions, though he believed that the two bison species should be lumped into ''Bison bison''. The

kouprey The kouprey (''Bos sauveli''), also known as forest ox is a forest-dwelling, wild bovine species native to Southeast Asia. A young male was sent to the Paris Zoological Park in 1937 and was described by the French zoologist Achille Urbain who ...

was not included in Simpson's taxonomy, while Bohlken (1958) considered the species to be a hybrid between banteng and cattle. Below is the Simpson (1945) taxonomy: * Subtribe Bovina (Gray, 1821) ** Genus ''Bibos'' (

Hodgson Hodgson is a surname. In United Kingdom, Britain, the Hodgson surname was the 173rd most common (766 per million) in 1881 and the 206th most common (650 per million) in 1998. In the United States, United States of America, Hodgson was the 3753rd mo ...

, 1837) *** ''Bibos javanicus'' (d’Alton, 1823) – Banteng *** ''Bibos gaurus'' (Hamilton-Smith, 1827) –

Gaur The gaur (''Bos gaurus''; ), also known as the Indian bison, is a bovine native to South Asia and Southeast Asia, and has been listed as Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 m ...

*** ''Bibos frontalis'' (Lambert, 1804) –

Gayal The gayal (''Bos frontalis''), also known as the Drung ox or mithun, is a large domestic cattle distributed in Northeast India, Bangladesh, Myanmar and in Yunnan, China.Simoons, F. J. (1984). ''Gayal or mithan''. In: Mason, I. L. (ed.) ''Evolut ...

** Genus '' Bison'' ( Hamilton-Smith, 1827) *** ''Bison bonasus'' (

Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...

, 1758) –

European bison The European bison (''Bison bonasus'') or the European wood bison, also known as the wisent ( or ), the zubr (), or sometimes colloquially as the European buffalo, is a European species of bison. It is one of two extant species of bison, along ...

*** ''Bison bison'' (

, 1758) –

American bison The American bison (''Bison bison'') is a species of bison native to North America. Sometimes colloquially referred to as American buffalo or simply Bubalina, buffalo (a different clade of bovine), it is one of two extant species of bison, alongs ...

** Genus ''

'' (

, 1758) *** Subgenus ''Poephagus'' (

Gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed o ...

, 1843) **** ''Bos mutus'' (Przewalski, 1883) – Wild yak **** ''Bos grunniens'' (Linnaeus, 1766) – Yak *** Subgenus ''

'' (

, 1758) **** †''Bos primigenius'' (Bojanus, 1827) – Aurochs **** ''Bos indicus'' (

, 1758) –

Zebu The zebu (; ''Bos indicus'' or ''Bos taurus indicus''), sometimes known in the plural as indicine cattle or humped cattle, is a species or subspecies of domestic cattle originating in the Indian sub-continent. Zebu are characterised by a fatty h ...

**** ''Bos taurus'' (

, 1758) –

Cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...

Subsequent taxonomic studies resulted ''Bibos'' to be reduced to as either a subgenus or a junior synonym of ''Bos''. As shown below: * Subtribe Bovina (Gray, 1821) ** Genus '' Bison'' ( Hamilton-Smith, 1827) *** ''Bison bonasus'' (Linnaeus, 1758) –

*** ''Bison bison'' (Linnaeus, 1758) –

** Genus ''

'' (Linnaeus, 1758) *** Subgenus ''Bibos'' (

, 1837) **** ''Bos javanicus'' (d’Alton, 1823) – Banteng **** ''Bos sauveli'' (Urbain, 1937) –

Kouprey The kouprey (''Bos sauveli''), also known as forest ox is a forest-dwelling, wild bovine species native to Southeast Asia. A young male was sent to the Paris Zoological Park in 1937 and was described by the French zoologist Achille Urbain who ...

**** ''Bos frontalis'' (Lambert, 1804) –

(including

gayal The gayal (''Bos frontalis''), also known as the Drung ox or mithun, is a large domestic cattle distributed in Northeast India, Bangladesh, Myanmar and in Yunnan, China.Simoons, F. J. (1984). ''Gayal or mithan''. In: Mason, I. L. (ed.) ''Evolut ...

) *** Subgenus ''

'' (Linnaeus, 1758) **** ''Bos taurus'' (Linnaeus, 1758) –

(including

and Aurochs) *** Subgenus ''Poephagus'' (

, 1843) **** ''Bos grunniens'' (Linnaeus, 1766) – Yak (including wild yak) However recent molecular work on mitochondrial DNA and the

Y-chromosome The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or abs ...

has completely revamped the evolutionary relationships among bovinans. These studies support of not only the inclusion of bison species into the genus ''Bos'', but offer two radically different positions for the

. According to the mitochondrial DNA, these studies support the

being closely related to the yak, while the

is more related to the aurochs. However a 2008 phylogenetic study using the Y-chromosome found the two bison species to form a clade. It also found that the yaks are an outgroup in relation to the rest of the bovinans, supporting their classification in the genus ''Poephagus''. Another study by Hassanin et al. (2013) using autosomal introns found support in the bison-yak clade. This suggests the mitochondrial genomes is result of

incomplete lineage sorting Incomplete lineage sorting, also termed hemiplasy, deep coalescence, retention of ancestral polymorphism, or trans-species polymorphism, describes a phenomenon in population genetics when ancestral gene copies fail to coalesce (looking backwards i ...

during

divergence In vector calculus, divergence is a vector operator that operates on a vector field, producing a scalar field giving the quantity of the vector field's source at each point. More technically, the divergence represents the volume density of t ...

of ''Bos'' and ''Bison'' from their common ancestors rather than further post- speciation gene flow (ancient hybridization between ''Bos'' and ''Bison''). There is evidence of limited gene flow from ''Bos primigenius taurus'' could account for the affiliation between wisent and cattle nuclear

genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...

s (in contrast to mitochondrial ones). These phylogenetic studies lead Groves and Grubb (2011), who conducted large scale taxonomic analysis on the world's ungulate species, to recommend classifying the two bison species as members of the genus ''Bos''. Below is the listing of species recognized by Groves and Grubb (2011) with species names following Castelló (2016) from ''Bovids of the World'': * Subtribe Bovina (Gray, 1821) ** Genus ''

'' (Linnaeus, 1758) *** ''Bos javanicus'' (d’Alton, 1823) – Banteng *** ''Bos sauveli'' (Urbain, 1937) –

*** ''Bos gaurus'' (Hamilton-Smith, 1827) –

*** ''Bos frontalis'' (Lambert, 1804) –

*** ''Bos mutus'' (Przewalski, 1883) – Wild yak *** ''Bos grunniens'' (Linnaeus, 1766) – Yak *** ''Bos bison'' (Linnaeus, 1758) –

*** ''Bos bonasus'' (Linnaeus, 1758) –

*** †''Bos caucasicus'' (Satunin, 1904) – Caucasian bison *** †''Bos primigenius'' (Bojanus, 1827) – Aurochs *** ''Bos indicus'' (Linnaeus, 1758) –

*** ''Bos taurus'' (Linnaeus, 1758) –

Fossil

The bovinans have a rich fossil record. According to the fossil record and molecular studies, Bubalina and Bovina diverged from one and another from a common ancestor around 13.7 million years ago in the Late Miocene. After arriving into Africa there was a rapid radiation of bovinan species in Africa in the Middle Pliocene. Among the diverse genera of African bovinans were two significant genera: ''

Pelorovis ''Pelorovis'' ("prodigious/monstrous sheep") is an extinct genus of African wild cattle which existed during the Pleistocene epoch.Alan Turner & Mauricio Anton: ''Evolving Eden, An Illustrated Guide to the Evolution of the African Large-Mammal Fau ...

'' and '' Leptobos''. According to anatomical and morphological study on the various species of ''Pelorovis'' and ''Leptobos'', it is believed the former genus evolved into ''

'' while the latter genus evolved into '' Bison'' during the

Late Pliocene Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...

of East Africa. Both lineages then left Africa and into Eurasia at the end of the Pliocene. While ''Bos'' inhabited much of Eurasia, some species of ''Bison'' had colonized North America by crossing over the

Bering Land Bridge Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 72 degrees north latitude in the Chukchi Sea; and on the south by the tip of ...

in two waves, the first being 135,000 to 195,000 years ago and the second being 21,000 to 45,000 years ago.An Alaska volcano and DNA reveal the timing of bison's arrival in North America
''

Alaska Dispatch News The ''Anchorage Daily News'' is a daily newspaper published by the Binkley Co., and based in Anchorage, Alaska. It is the most widely read newspaper and news website (adn.com) in the state of Alaska. The newspaper is headquartered in Anchorage, ...

'', Yereth Rosen, March 27, 2017. Retrieved 28 March 2017. The exact relationships between fossil and extant bovinans problematic. Below is the list of fossil species that have been described so far (listed alphabetically): * Subtribe Bovina (Gray, 1821) ** Genus †'' Adjiderebos'' (Dubrovo & Burchak-Abramovich, 1984) *** †''Adjiderebos cantabilis'' (Dubrovo & Burchak-Abramovich, 1984) ** Genus '' Bison'' (Hamilton-Smith, 1827) *** †''

Bison antiquus ''Bison antiquus'', the antique bison or ancient bison, is an extinct species of bison that lived in Late Pleistocene North America until around 10,000 years ago. It was one of the most common large herbivores on the North American continent dur ...

'' (Leidy, 1852) *** †''Bison georgicus'' (Burchak-Abramovich & Vekua, 1994) *** †''Bison hanaizumiensis'' (Matsumoto & Mori, 1956) *** †''

Bison latifrons ''Bison latifrons'', also known as the giant bison or long-horned bison, is an extinct species of bison that lived in North America during the Pleistocene epoch ranging from Alaska to Mexico. It was the largest and heaviest bovid ever to live in ...

'' (Harlan, 1825) *** †''Bison menneri'' (Sher, 1997) *** †'' Bison palaeosinensis'' (Teilhard & Piveteau, 1930) *** †'' Bison priscus'' (Bojanus, 1827) *** †'' Bison schoetensacki'' (Freudenberg, 1914) *** †''Bison sivalensis'' (Falconer, 1878) *** †''Bison tamanensis'' (Vereshchagin, 1959) *** †''Bison voigtstedtensis'' (Fischer, 1965) ** Genus ''

'' (Linnaeus, 1758) *** Subgenus ''Bos'' (Linnaeus, 1758) **** †''Bos acutifrons'' (Lydekker, 1877) **** †''Bos buiaensis'' (Martínez-Navarro ''et al''., 2009) **** †''Bos caucasicus'' (Burchak-Abramovich & Vekua, 1980) **** †'' Bos planifrons'' (Lydekker, 1877) *** Subgenus ''Bibos'' **** †'' Bos palaesondaicus'' (Dubois, 1908) *** Subgenus ''Poephagus'' (Gray, 1843) **** †''Bos baikalensis'' (Verestchagin, 1954) ** Genus †'' Epileptobos'' (Hooijer, 1956) *** †''Epileptobos groeneveldtii'' (Dubois, 1908) ** Genus †'' Ioribos'' (Vekua, 1972) *** †''Ioribos aceros'' (Vekua, 1972) ** Genus †'' Leptobos'' (Rütimeyer, 1877) *** Subgenus †''Leptobos'' (Rütimeyer, 1877) **** †''Leptobos elatus'' (Croizet & Pomel, 1853) **** †''Leptobos falconeri'' (Rütimeyer, 1877) **** †''Leptobos furtivus'' (Duvernois & Guérin, 1989) *** Subgenus †''Smertiobos'' (Duvernois, 1992) **** †''Leptobos bravardi'' (Duvernois, 1989) **** †''Leptobos brevicornis'' (Hu & Qi, 1975) **** †''Leptobos crassus'' (Jia & Wang, 1978) **** †''Leptobos etruscus'' (Falconer, 1859) ** Genus †''

'' (Reck, 1928) *** †''Pelorovis howelli'' (Hadjouis & Sahnouni, 2006) *** †''Pelorovis kaisensis'' (Geraads & Thomas, 1994) *** †''Pelorovis oldowayensis'' (Reck, 1928) *** †''Pelorovis praeafricanus'' (Arambourg, 1979) *** †''Pelorovis turkanensis'' (Harris, 1991) ** Genus †'' Platycerabos'' (Barbour & Schultz, 1942) *** †''Platycerabos dodsoni'' (Barbour & Schultz, 1941) ** Genus †'' Protobison'' (Burchak-Abramovich, Gadzhiev & Vekua, 1980) *** †''Protobison kushkunensis'' (Burchak-Abramovich, Gadzhiev & Vekua, 1980) ** Genus †'' Urmiabos'' (Burchak-Abramovich, 1950) *** †''Urmiabos azerbaidzanicus'' (Burchak-Abramovich, 1950) ** Genus †'' Yakopsis'' (Kretzoi, 1954) *** †''Yakopsis stenometopon'' (Rütimeyer, 1865)

References

{{Taxonbar, from=Q16533792 Bovines Extant Miocene first appearances Animal subtribes Taxa named by John Edward Gray