View source for Bovidae

{{Short description|Family of mammals belonging to even-toed ungulates}}
{{Good article}}
{{Automatic taxobox
| fossil_range = {{Fossil range|20|0}}<small>Early [[Miocene]]&nbsp;[[Holocene|present]]</small>
| image = Bovidae-0001.jpg
| image_caption = Example Bovidae (clockwise from top left) – [[addax]] (''Addax nasomaculatus''), [[Bos taurus|domestic cattle]] (''Bos taurus''), [[mountain gazelle]] (''Gazella gazella''), [[impala]] (''Aepyceros melampus''), [[blue wildebeest]] (''Connochaetes taurinus''), and [[mouflon]] (''Ovis gmelini'')
| display_parents = 2
| taxon = Bovidae
| authority = [[John Edward Gray|Gray]], 1821
| type_genus = ''[[Bos]]''
| type_genus_authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]
| subdivision_ranks = Subfamilies
| subdivision = *[[Aepycerotinae]] (1 genus)<br />
* [[Alcelaphinae]] (4 genera)
* [[Antilopinae]] (3 tribes and 14 genera)
* [[Bovinae]] (3 tribes and 15 genera)
* [[Caprinae]] (3 tribes and 15 genera)
* [[Cephalophinae]] (3 genera)
* [[Hippotraginae]] (3 genera)
* [[Nesotragus|Nesotraginae]] (1 genus)
* [[Oreotraginae]] (1 genus)
* [[Reduncinae]] (3 genera)
* †[[Hypsodontinae]]
Alternate taxonomy:
* [[Antilopinae]] (10 tribes and 45 genera)
* [[Bovinae]] (3 tribes and 15 genera)
* †[[Hypsodontinae]]
}}
The '''Bovidae''' comprise<!-- The use "are" replace "comprise" bovid have muskoxen and oxen--> the [[family (biology)|biological family]] of [[cloven-hoof]]ed, [[ruminant]] [[mammal]]s that includes [[Bos|cattle]], [[bison]], [[Bubalina|buffalo]], [[antelope]]s (including [[Caprinae|goat-antelopes]]), [[Ovis|sheep]] and [[Capra (genus)|goats]].  A member of this family is called a '''bovid'''. With 143 [[Extant taxon|extant]] species and 300 known [[Extinction|extinct species]], the family Bovidae consists of 11 (or two) major [[subfamilies]] and thirteen major tribes. The family evolved 20 million years ago, in the early [[Miocene]].
The bovids show great variation in size and [[Fur|pelage]] colouration. Except some [[Polled livestock|domesticated forms]], all male bovids have two or more [[horn (anatomy)|horns]], and in many species, females possess horns, too. The size and shape of the horns vary greatly, but the basic structure is always one or more pairs of simple bony protrusions without branches, often having a spiral, twisted or fluted form, each covered in a permanent sheath of [[keratin]]. Most bovids bear 30 to 32 teeth.
Most bovids are [[diurnality|diurnal]]. Social activity and feeding usually peak during dawn and dusk. Bovids typically rest before dawn, during midday, and after dark. They have various methods of social organisation and [[social behaviour]], which are classified into solitary and gregarious behaviour. Bovids use different forms of vocal, olfactory, and tangible communication. Most species alternately feed and ruminate throughout the day. While small bovids forage in dense and closed habitat, larger species feed on high-fiber vegetation in open grasslands. Most bovids are [[Polygynous mating systems|polygynous]]. <!--'Mature bovids mate at least once a year and smaller species may even mate twice.' This doesn't tell us anything really meaningful! Do any large species mate more than twice a year? If not, it is just saying that some species, whatever their size, mate once, others twice. Which is more prevalent? Is twice a year more prevalent among smaller species? Do immature specimens even mate?-->Mature bovids mate at least once a year and smaller species may even mate twice. In some species, neonate bovids remain hidden for a week to two months, regularly nursed by their mothers; in other species, neonates are followers, accompanying their dams, rather than tending to remain hidden.
The greatest diversities of bovids occur in [[Africa]]. The maximum concentration of species is in the [[savanna]]s of [[Eastern Africa]]. Other bovid species also occur in Europe, Asia, and North America. Bovidae includes a number of [[Domestication|domesticated]] species, including three whose use has spread worldwide, namely [[cattle]], [[sheep]], and [[goats]]. [[Dairy products]], such as [[milk]], [[butter]], and [[cheese]], are manufactured largely from domestic cattle. Bovids are also raised for their [[leather]], [[meat]], and [[wool]].
==Naming and etymology==
The name "Bovidae" was given by the British zoologist [[John Edward Gray]] in 1821.<ref name=MSW3>{{MSW3 Artiodactyla |id=14200485 |heading=Family Bovidae}}</ref> The word "Bovidae" is the combination of the prefix ''bov-'' (originating from [[Latin]] ''bos'', "ox", through [[Late Latin]] ''bovinus'') and the suffix ''-idae''.<ref name=mw>{{cite web|title=Bovidae|url=http://www.merriam-webster.com/dictionary/bovidae|website=Merriam-Webster online dictionary|access-date=7 October 2014|archive-date=11 October 2014|archive-url=https://web.archive.org/web/20141011131843/http://www.merriam-webster.com/dictionary/bovidae|url-status=live}}</ref>
==Taxonomy==
{{main|List of bovids}}
The [[Family (biology)|family]] Bovidae is placed in the [[Order (biology)|order]] [[Artiodactyla]] (which includes the even-toed [[ungulate]]s). It includes 143 extant species, accounting for nearly 55% of the [[ungulate]]s, and 300 known extinct species.<ref name=adw>{{cite web |last1=Gomez |first1=W. |last2=Patterson |first2=T. A. |last3=Swinton |first3=J. |last4=Berini |first4=J. |title=Bovidae: antelopes, cattle, gazelles, goats, sheep, and relatives |url=http://animaldiversity.ummz.umich.edu/accounts/Bovidae/ |website=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |access-date=7 October 2014 |archive-date=7 October 2014 |archive-url=https://web.archive.org/web/20141007065046/http://animaldiversity.ummz.umich.edu/accounts/Bovidae/ |url-status=live }}</ref>
Until the beginning of the 21st century it was understood that the family [[Moschidae]] (musk deer) was [[sister taxon|sister]] to [[Cervidae]]. However, a 2003 [[phylogenetic]] study by Alexandre Hassanin (of [[National Museum of Natural History (France)|National Museum of Natural History, France]]) and colleagues, based on [[mitochondria]]l and [[nucleus (biology)|nuclear]] analyses, revealed that [[Moschidae]] and Bovidae form a [[clade]] sister to [[Cervidae]]. According to the study, Cervidae [[genetic divergence|diverged]] from the Bovidae-Moschidae clade 27 to 28 million years ago.<ref name="Hassanin2003">{{cite journal | last1=Hassanin | first1=A. | last2=Douzery | first2=E. J. P. | title=Molecular and morphological phylogenies of Ruminantia and the alternative position of the Moschidae | journal=Systematic Biology | date=2003 | volume=52 | issue=2 | pages=206–28 | doi=10.1080/10635150390192726 | url=https://www.researchgate.net/publication/10760976 | pmid=12746147| doi-access=free }}</ref> The following [[cladogram]] is based on the 2003 study.<ref name="Hassanin2003"/>
{{Clade
| style=font-size: 100%; line-height:100%
| label1= [[Ruminantia]]
| 1={{clade
| label1= [[Tragulina]]
| 1={{clade
| label1=
| 1=[[Tragulidae]] [[File:Tragulus napu - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|50 px]]
}}
| label2= [[Pecora]]
| 2={{clade
| label1=
| 1=[[Antilocapridae]] [[File:Antilocapra white background.jpg|50 px]]
| 2=[[Giraffidae]] [[File:Giraffa camelopardalis Brockhaus white background.jpg|50 px]]
| 3={{clade
| label1=
| 1=[[Cervidae]] [[File:The deer of all lands (1898) Hangul white background.png|50 px]]
| 2={{clade
| label1=
| 1=[[Moschidae]] [[File:Moschus chrysogaster white background.jpg|50 px]]
| 2='''Bovidae''' [[File:Birds and nature (1901) (14562088237) white background.jpg |50px]]
}}
}}
}}
}}
}}
Molecular studies have supported [[monophyly]] in the family Bovidae (a group of organisms comprises an ancestral species and all their descendants).<ref name=gatesy>{{cite journal |last1=Gatesy |first1=J. |last2=Amato |first2=G. |last3=Vrba |first3=E. |last4=Schaller|first4=G. |title=A cladistic analysis of mitochondrial ribosomal DNA from the Bovidae |journal=Molecular Phylogenetics and Evolution|volume=7|issue=3|year=1997|pages=303–19 |doi=10.1006/mpev.1997.0402|pmid=9187090 |doi-access=free |bibcode=1997MolPE...7..303G }}</ref><ref name=vrba2005>{{cite journal|last1=Fernández |first1=M. H.|last2=Vrba |first2=E. S. |title=A complete estimate of the phylogenetic relationships in Ruminantia: a dated species-level supertree of the extant ruminants |journal=Biological Reviews |date=2005 |volume=80 |issue=2 |pages=269–302 |doi=10.1017/S1464793104006670 |pmid=15921052|s2cid=29939520}}</ref> The number of [[subfamily|subfamilies]] in Bovidae is disputed, with suggestions of as many as ten and as few as two subfamilies.<ref name=vrba2005/> However, molecular, [[Morphological (biology)|morphological]] and fossil evidence indicates the existence of eight distinct subfamilies: [[Aepycerotinae]] (consisting of just the impala), [[Alcelaphinae]] (bontebok, hartebeest, wildebeest and relatives), [[Antilopinae]] (several antelopes, gazelles, and relatives), [[Bovinae]] (cattle, buffaloes, bison and other antelopes), [[Caprinae]] (goats, sheep, ibex, serows and relatives), [[Cephalophinae]] (duikers), [[Hippotraginae]] (addax, oryx and relatives) and [[Reduncinae]] (reedbuck and kob antelopes). In addition, three extinct subfamilies are known: [[Hypsodontinae]] (mid-[[Miocene]]), [[Oiocerinae]] ([[Turolian]]) and the subfamily [[Tethytraginae]], which contains ''[[Tethytragus]]'' (mid-[[Miocene]]).<ref name=harrison2>{{cite book |last1=Harrison |first1=T. |title=Paleontology and Geology of Laetoli Human Evolution in Context |date=2011 |publisher=Springer |location=Dordrecht|isbn=978-9048-199-624 |pages=363–465}}</ref><ref name=tethytragus>{{cite journal |last1=Demiguel |first1=D. |last2=Sánchez |first2=I. M. |last3=Alba |first3=D. M. |last4=Galindo |first4=J. |last5=Robles |first5=J. M. |last6=Moyà-Solà |first6=S. |title=First evidence of ''Tethytragus'' Azanza and Morales, 1994 (Ruminantia, Bovidae), in the Miocene of the Vallès-Penedès Basin (Spain) |journal=Journal of Vertebrate Paleontology |volume=32 |issue=6 |pages=1457–62 |doi=10.1080/02724634.2012.696082|year=2012 |bibcode=2012JVPal..32.1457D |s2cid=86664298 }}</ref>
In 1992, Alan W . Gentry of the [[Natural History Museum, London]] divided the eight major subfamilies of Bovidae into two major clades on the basis of their evolutionary history: the Boodontia, which comprised only the Bovinae, and the Aegodontia, which consisted of the rest of the subfamilies. Boodonts have somewhat [[Primitive (phylogenetics)|primitive]] teeth, resembling those of [[oxen]], whereas aegodonts have more [[Evolutionary progress|advanced]] teeth like those of goats.<ref name=harrison>{{cite book |last1=Harrison |first1=T. |title=Neogene Paleontology of the Manonga Valley, Tanzania: A Window into the Evolutionary History of East Africa |date=1997 |publisher=Plenum Press |location=New York |isbn=978-0-306-45471-4 |page=113}}</ref>
A controversy exists about the recognition of [[Peleinae]] and [[Pantholopinae]], comprising the genera ''[[Grey rhebok|Pelea]]'' and ''[[Pantholops]]'' respectively, as [[subfamily|subfamilies]]. In 2000, American biologist [[George Schaller]] and palaeontologist [[Elisabeth Vrba]] suggested the inclusion of ''[[Grey rhebok|Pelea]]'' in [[Reduncinae]],<ref name=vrba2000>{{cite book|last1=Vrba |first1=E. S. |last2=Schaller |first2=G.|title=Antelopes, Deer, and Relatives: Fossil Record, Behavioral Ecology, Systematics, and Conservation |date=2000 |publisher=Yale University Press |location=New Haven |isbn=978-0300-081-428}}</ref> though the grey rhebok, the sole species of ''[[Grey rhebok|Pelea]]'', is highly different from kobs and reduncines in morphology.<ref name=msw3(1)>{{MSW3_Artiodactyla |page=719|id = 14200959}}</ref> ''[[Pantholops]]'', earlier classified in the [[Antilopinae]], was later placed in its own subfamily, [[Pantholopinae]]. However, molecular and morphological analysis supports the inclusion of ''[[Pantholops]]'' in [[Caprinae]].<ref name=msw3(2)>{{MSW3_Artiodactyla |page=699|id = 14200851}}</ref>
Below is a cladogram based on Yang ''et al''.'','' 2013 and Calamari, 2021:<ref>{{Cite journal|last1=Yang|first1=Chengzhong|last2=Xiang|first2=Changkui|last3=Qi|first3=Wenhua|last4=Xia|first4=Shan|last5=Tu|first5=Feiyun|last6=Zhang|first6=Xiuyue|last7=Moermond|first7=Timothy|last8=Yue|first8=Bisong|date=2013-06-01|title=Phylogenetic analyses and improved resolution of the family Bovidae based on complete mitochondrial genomes|url=https://www.sciencedirect.com/science/article/pii/S0305197812002670|journal=Biochemical Systematics and Ecology|language=en|volume=48|pages=136–143|doi=10.1016/j.bse.2012.12.005|bibcode=2013BioSE..48..136Y |issn=0305-1978|access-date=2022-01-31|archive-date=2022-01-31|archive-url=https://web.archive.org/web/20220131032745/https://www.sciencedirect.com/science/article/pii/S0305197812002670|url-status=live}}</ref><ref name=":0">{{Cite journal|last1=Matthee|first1=Conrad A.|last2=Robinson|first2=Terence J.|date=1999-06-01|title=CytochromebPhylogeny of the Family Bovidae: Resolution within the Alcelaphini, Antilopini, Neotragini, and Tragelaphini|url=https://www.sciencedirect.com/science/article/pii/S1055790398905735|journal=Molecular Phylogenetics and Evolution|language=en|volume=12|issue=1|pages=31–46|doi=10.1006/mpev.1998.0573|pmid=10222159 |bibcode=1999MolPE..12...31M |issn=1055-7903}}</ref><ref name=":1">{{Cite journal|last=Calamari|first=Zachary T.|date=June 2021|title=Total Evidence Phylogenetic Analysis Supports New Morphological Synapomorphies for Bovidae (Mammalia, Artiodactyla)|url=https://bioone.org/journals/american-museum-novitates/volume-2021/issue-3970/3970.1/Total-Evidence-Phylogenetic-Analysis-Supports-New-Morphological-Synapomorphies-for-Bovidae/10.1206/3970.1.full|journal=American Museum Novitates|issue=3970|pages=1–38|doi=10.1206/3970.1|s2cid=235441087|issn=0003-0082|access-date=2022-02-08|archive-date=2023-03-14|archive-url=https://web.archive.org/web/20230314124448/https://bioone.org/journals/american-museum-novitates/volume-2021/issue-3970/3970.1/Total-Evidence-Phylogenetic-Analysis-Supports-New-Morphological-Synapomorphies-for-Bovidae/10.1206/3970.1.short|url-status=live|hdl=2246/7267|hdl-access=free}}</ref>
{{clade
|style=font-size:90%;line-height:100%;width:700px;
|label1=  '''Bovidae'''
|1={{clade
|label1=  Boodontia
|sublabel1=  ([[Bovinae]])
|1={{clade
|label1=
|1={{clade
|label1=
|1=[[Bovini]] (bison, buffalo, cattle, etc.) [[File:Birds and nature (1901) (14562088237) white background.jpg|50 px]]
}}
|2={{clade
|label1=
|1=[[Boselaphini]] (nilgai and four-horned antelope) [[File:Haeckel Antilopina Tetracerus quadricornis.jpg|50 px]]
|2=[[Tragelaphini]] (kudus, nyalas etc.) [[File:The book of antelopes (1894) Tragelaphus angasi white background.png|50 px]]
}}
}}
|label2= Aegodontia
|2={{clade
|label1=
|1={{clade
|label1=
|1=[[Aepycerotinae]] (impala) [[File:The book of antelopes (1894) Aepyceros melampus white backround.png|50 px]]
|2=[[Nesotragus|Nesotraginae]] (suni and bates's antelope)
}}
|2={{clade
|label1=
|1={{clade
|label1=
|1=[[Antilopinae]] (gazelles, springbok, dik-dik, royal antelope, saiga, etc.) [[File:The book of antelopes (1894) Gazella thomsoni white background.png|50 px]]
|2={{clade
|label1=
|1=[[Cephalophinae]] (duikers etc.) [[File:The book of antelopes (1894) Cephalophus natalensis white background.png|50 px]]
|2=[[Oreotraginae]] (klipspringer)
}}
}}
|2={{clade
|label1=
|1=[[Reduncinae]] (kobs, reedbucks, waterbucks etc.) [[File:The book of antelopes (1894) Cobus defassa white background.png|50 px]]
|2={{clade
|label1=
|1=[[Caprinae]] (chamois, sheep, ibexes, goats, muskox, etc.) [[File:Walia ibex illustration white background.png|50 px]]
|2={{clade
|label1=
|1=[[Alcelaphinae]] (hartebeest, topi, wildebeest etc.) [[File:The book of antelopes (1894) Bubalis busephalus white backround.png|50 px]]
|2=[[Hippotraginae]] (sable antelopes, oryxes etc.) [[File:The book of antelopes (1894) Hippotragus niger I white background.png|50 px]]
}}
}}
}}
}}
}}
}}
}}
Alternatively, all members of the Aegodontia, can be classified within the subfamily Antilopinae, with the individual subfamilies being tribes in this treatment.<ref name=":0" /><ref name=":1" />
==Evolutionary history==
===Early Miocene and before===
[[File:Eotragus sansaniensis.JPG|thumbnail|Skull of ''Eotragus sansaniensis'', a species of the ancient bovid genus ''[[Eotragus]]''
]]
In the early Miocene, bovids began diverging from the [[cervid]]s (deer) and [[giraffid]]s. The earliest bovids, whose presence in Africa and Eurasia in the latter part of early Miocene (20 [[Million years ago|Mya]]) has been ascertained, were small animals, somewhat similar to modern [[gazelle]]s, and probably lived in [[woodland]] environments.<ref name=MamEv/> ''[[Eotragus]]'', the earliest known bovid, weighed {{convert|18|kg|lb|abbr=on}} and was nearly the same in size as the [[Thomson's gazelle]].<ref name=prothero/> Early in their evolutionary history, the bovids split into two main clades: Boodontia (of Eurasian origin) and Aegodontia (of African origin). This early [[Genetic divergence|split]] between Boodontia  and Aegodontia has been attributed to the continental divide between these land masses. When these continents were later rejoined, this barrier was removed, and both groups expanded into the territory of the other.<ref name=Hassanin>{{cite journal | author = Hassanin, D. | year = 1999 | title = The tribal radiation of the family Bovidae (Artiodactyla) and the evolution of the mitochondrial cytochrome ''b'' gene | journal = [[Molecular Phylogenetics and Evolution]] | volume = 13 | issue = 2 | pages = 227–43 | url = http://www.isem.cnrs.fr/IMG/pdf/Hassanin_1999-MPE.pdf | archive-url = https://web.archive.org/web/20110720212305/http://www.isem.cnrs.fr/IMG/pdf/Hassanin_1999-MPE.pdf | archive-date = 2011-07-20 | pmid = 10603253 | doi = 10.1006/mpev.1999.0619 | last2 = Douzery | first2 = E.J. | bibcode = 1999MolPE..13..227H }}</ref> The tribes [[Bovini]] and Tragelaphini diverged in the early Miocene.<ref name=gilbert/> Bovids are known to have reached the [[Americas]] in the Pleistocene by crossing the [[Bering land bridge]].<ref name=prothero/>
The present genera of Alcelaphinae appeared in the [[Pliocene]]. The extinct Alcelaphine genus ''Paramularius'', which was the same in size as the hartebeest, is believed to have come into being in the Pliocene, but became extinct in the middle [[Pleistocene]].<ref name=vrba2005/> Several genera of Hippotraginae are known since the Pliocene and Pleistocene. This subfamily appears to have diverged from the Alcelaphinae in the latter part of early Miocene.<ref name=gilbert/> The Bovinae are believed to have [[genetic divergence|diverged]] from the rest of the Bovidae in the early [[Miocene]].<ref name=Bibi2013>{{cite journal |author=Bibi, F. | date = August 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 | page = 166 | doi = 10.1186/1471-2148-13-16| pmid = 23339550 | pmc = 3607928 | doi-access = free }}</ref> The Boselaphini became extinct in Africa in the early Pliocene; their latest fossils were excavated in [[Langebaanweg]] (South Africa) and [[Lothagam]] (Kenya).<ref name=geraads>{{cite journal|last1=Geraads|first1=D.|last2=El Boughabi|first2=S.|last3=Zouhri|first3=S.|title=A new caprin bovid (Mammalia) from the late Miocene of Morocco|journal=Palaeontologica Africana|date=2012|issue=47|pages=19–24|issn=0078-8554}}</ref>
===Middle Miocene===
The middle Miocene marked the spread of the bovids into China and the Indian subcontinent.<ref name=prothero/> According to Vrba, the [[Radiation (biology)|radiation]] of the subfamily Alcelaphinae began in the latter part of middle Miocene.<ref name=vrba2005/> The Caprinae tribes probably diverged in the early middle Miocene. The Caprini emerged in the middle Miocene, and seem to have been replaced by other bovids and cervids in Eurasia.<ref name=kingdon1989>{{cite book|last1=Kingdon|first1=J.|title=East African Mammals: An Atlas of Evolution in Africa (Volume III, Part C)|date=1989|publisher=University of Chicago press|location=Chicago|isbn=978-0-226-43724-8|pages=1–33}}</ref> The earliest fossils of the antilopines are from the middle Miocene, though studies show the existence of the subfamily from the early Miocene. [[Speciation]] occurred in the [[Tribe (biology)|tribe]] [[Antilopini]] during the middle or upper Miocene, mainly in Eurasia. Tribe [[Neotragini]] seems to have appeared in Africa by the end of Miocene, and had become widespread by the Pliocene.<ref name=gilbert>{{cite book|last1=Gilbert|first1=W. H.|last2=Asfaw|first2=B.|title=Homo Erectus: Pleistocene Evidence from the Middle Awash, Ethiopia|date=2008|publisher=University of California Press|location=Berkeley|isbn=978-0-520-25120-5|pages=45–84}}</ref>
===Late Miocene===
By the late Miocene, around 10 Mya, the bovids rapidly [[Genetic diversity|diversified]], leading to the creation of 70 new genera.<ref name=prothero>{{cite book|last1=Prothero|first1=D. R.|last2=Schoch|first2=R. M.|title=Horns, Tusks, and Flippers: the Evolution of Hoofed Mammals|date=2002|publisher=Johns Hopkins University Press|location=Baltimore|isbn=978-0-8018-7135-1|pages=87–90}}</ref> This late Miocene radiation was partly because many bovids became adapted to more open, grassland habitats.<ref name=MamEv>{{cite book |author1=Savage, R.J.G. |author2=Long, M.R. |year=1986 |title= Mammal Evolution: an illustrated guide|url=https://archive.org/details/mammalevolutioni0000sava |url-access=registration |publisher= Facts on File|location=New York|pages= [https://archive.org/details/mammalevolutioni0000sava/page/232 232–5]|isbn= 978-0-8160-1194-0}}</ref> The Aepycerotinae first appeared in the late Miocene, and no significant difference in the sizes of the primitive and modern impala has been noted.<ref name=vrba1984>{{cite book|last1=Stanley|first1=S. M.|last2=Eldredge|first2=N.|title=Living Fossils|date=1984|publisher=Springer|isbn=978-1461-382-737|pages=62–79|chapter=Evolutionary Pattern and Process in the Sister-Group Alcelaphini-Aepycerotini (Mammalia: Bovidae)}}</ref> Fossils of ovibovines, a tribe of Caprinae, in Africa date back to the late Miocene.<ref name=gilbert/> The earliest Hippotragine fossils date back to the late Miocene, and were excavated from sites such as Lothagam and [[Awash Valley]].<ref name=gilbert/> The first African fossils of Reduncinae date back to 6-7 Mya.<ref name=vrba1995>{{cite book|last1=Vrba|first1=E. S.|last2=Burckle|first2=L. H.|last3=Partridge|first3=T. C.|last4=Denton|first4=G. H.|title=Paleoclimate and Evolution, with Emphasis on Human Origins|date=1995|publisher=Yale University Press|location=New Haven|isbn=978-0300-063-486|pages=24–5}}</ref> Reduncinae and Peleinae probably diverged in the mid-Miocene.<ref name=vrba2005/>
==Characteristics==
[[File:Serengeti Bueffel2.jpg|thumbnail|Bovids have unbranched horns.]]
All bovids have the similar basic form - a snout with a blunt end, one or more pairs of horns (generally present on males) immediately after the oval or pointed ears, a distinct neck and limbs, and a tail varying in length and bushiness among the species.<ref name="walton">{{cite book |last1=Walton |first1=D.W. |title=Fauna of Australia (Volume 1B) |date=1989 |publisher=Australian Government Publication Service |location=Canberra |isbn=978-0644-060-561 |pages=1–14}}</ref> Most bovids exhibit [[sexual dimorphism]], with males usually larger as well as heavier than females. Sexual dimorphism is more prominent in medium- to large-sized bovids. All bovids have four toes on each foot – they walk on the central two (the [[hoof|hooves]]), while the outer two (the [[dewclaw]]s) are much smaller and rarely touch the ground.<ref name="adw" />
The bovids show great variation in size: the [[gaur]] can weigh more than {{convert|1500|kg|lb|abbr=on}}, and stand {{convert|2.2|m|in|abbr=on}} high at the shoulder.<ref>{{cite book |author=Hassanin, A. |chapter=Systematics and Phylogeny of Cattle |pages=1–18 |date=2015 |title=The Genetics of Cattle |edition=Second |editor1=Garrick, D. |editor2=Ruvinsky, A. |publisher=Cabi |location=Oxfordshire, Boston |isbn=978-1-78064-221-5 |chapter-url=https://books.google.com/books?id=RYgXBgAAQBAJ&pg=PA7 |access-date=2021-01-13 |archive-date=2023-03-14 |archive-url=https://web.archive.org/web/20230314124448/https://books.google.com/books?id=RYgXBgAAQBAJ&pg=PA7 |url-status=live }}</ref> The [[royal antelope]], in sharp contrast, is only {{convert|25|cm|in|abbr=on}} tall and weighs at most {{convert|3|kg|lb|abbr=on}}.<ref name="royal">{{cite web |last1=Huffman |first1=B. |title=Royal antelope |url=http://www.ultimateungulate.com/Artiodactyla/Neotragus_pygmaeus.html |website=Ultimate Ungulate |access-date=8 October 2014 |archive-url=https://web.archive.org/web/20141216022820/http://www.ultimateungulate.com/Artiodactyla/Neotragus_pygmaeus.html |archive-date=16 December 2014 }}</ref> The [[klipspringer]], another small antelope, stands {{convert|45|-|60|cm|in|abbr=on}} at the shoulder and weighs just {{convert|10|-|20|kg|lb|abbr=on}}.<ref name="klipspringer">{{cite book |last1=Hildyard |first1=A. |title=Endangered Wildlife and Plants of the World |date=2001 |publisher=Marshall Cavendish |location=New York |isbn=978-0-7614-7200-1 |pages=769–70}}</ref>
Differences occur in [[Coat (animal)|pelage]] colouration, ranging from a pale white (as in the [[Arabian oryx]])<ref name="oryx">{{EOL|331080| Oryx leucoryx}}</ref> to black (as in the [[black wildebeest]]).<ref name="blackwildebeest">{{cite web |url=http://animaldiversity.ummz.umich.edu/accounts/Connochaetes_gnou/ |title=''Connochaetes gnou'': black wildebeest |author1=Lundrigan, B. |author2=Bidlingmeyer, J. |year=2000 |work=Animal Diversity Web |publisher=University of Michigan |access-date=2013-08-21 |archive-date=2013-10-05 |archive-url=https://web.archive.org/web/20131005033834/http://animaldiversity.ummz.umich.edu/accounts/Connochaetes_gnou/ |url-status=live }}</ref> However, only the intermediate shades, such as brown and reddish brown (as in the [[reedbuck]]), are commonly observed.<ref name="estes">{{cite book |last1=Estes |first1=R. D. |title=The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates |date=2004 |publisher=University of California Press |location=Berkeley |isbn=978-0-520-08085-0 |pages=[https://archive.org/details/isbn_0520080858/page/7 7–25] |edition=4th |url=https://archive.org/details/isbn_0520080858/page/7 }}</ref> In several species, females and juveniles exhibit a light-coloured coat, while those of males darken with age. As in the [[wildebeest]], the coat may be marked with prominent or faint stripes. In some species such as the [[addax]], the coat colour can vary by the season.<ref name="Krausman2007">{{cite journal |author1=Krausman, P.R. |author2=Casey, A.L. |year=2012 |title=''Addax nasomaculatus'' |journal=Mammalian Species |pages=1–4 |doi=10.1644/807.1 |issue=807|s2cid=198123785 |doi-access= }}</ref> [[Scent gland]]s and [[sebaceous gland]]s are often present.<ref name="walton" />
[[File:Oryx gazella male 8054 b.jpg|thumb|left|The [[gemsbok]] has conspicuous markings on its face, which conceal the eye, and on its legs. These may have a role in communication.<ref name="Caro" />]]
Some species, such as the [[gemsbok]], [[sable antelope]], and [[Grant's gazelle]], are [[camouflage]]d with strongly [[disruptive coloration|disruptive]] facial markings that conceal the highly recognisable eye.<ref>{{cite book |author=Cott, H. B. |author-link=Hugh B. Cott |year=1940 |title=Adaptive Coloration in Animals |publisher=Methuen |location=London |pages=88 and plate 25|title-link=Adaptive Coloration in Animals }}</ref> Many species, such as gazelles, may be made to look flat, and hence to blend into the background, by [[countershading]].<ref name="Kiltie">{{cite journal |title=Countershading: Universally deceptive or deceptively universal? |author=Kiltie, R.A. |journal=Trends in Ecology & Evolution |date=January 1998 |volume=3 |issue=1 |pages=21–23 |doi=10.1016/0169-5347(88)90079-1|pmid=21227055 }}</ref> The outlines of many bovids are broken up with bold disruptive colouration, the strongly contrasting patterns helping to delay recognition by predators.<ref>{{cite book |author=Cott, H. B. |author-link=Hugh B. Cott |year=1940 |title=Adaptive Coloration in Animals |publisher=Methuen |location=London |page=[https://archive.org/details/adaptivecolorati00cott/page/53 53]|title-link=Adaptive Coloration in Animals }}</ref> However, all the Hippotraginae (including the gemsbok) have pale bodies and faces with conspicuous markings. The zoologist [[Tim Caro]] describes this as difficult to explain, but given that the species are diurnal, he suggests that the markings may function in communication. Strongly contrasting leg colouration is common only in the Bovidae, where for example ''Bos'', ''Ovis'', [[bontebok]] and gemsbok have white stockings. Again, communication is the likely function.<ref name="Caro">{{cite journal |last1=Caro |first1=Tim |author1-link=Tim Caro |title=Contrasting colouration in terrestrial mammals |journal=Philosophical Transactions of the Royal Society B |date=2009 |volume=364 |issue=1516 |pages=537–548 |doi=10.1098/rstb.2008.0221 |pmc=2674080 |pmid=18990666 }}</ref>
Excepting some domesticated forms, all male bovids have horns, and in many species, females, too, possess horns. The size and shape of the horns vary greatly, but the basic structure is a pair of simple bony protrusions without branches, often having a spiral, twisted, or fluted form, each covered in a permanent sheath of keratin. Although horns occur in a single pair on almost all bovid species, there are exceptions such as the [[four-horned antelope]]<ref name="leslie">{{cite journal |last1=Leslie |first1=D. M. |last2=Sharma |first2=K. |title=Tetracerus quadricornis (Artiodactyla: Bovidae) |journal=Mammalian Species |date=25 September 2009 |issue=843 |pages=1–11 |doi=10.1644/843.1|doi-access=free }}</ref> and the [[Jacob sheep]].<ref name="jacobsheep">{{cite book |last1=E.C. |first1=Mungall |title=Exotic Animal Field Guide: Nonnative Hoofed Mammals in the United States |date=2007 |publisher=Texas A&M University Press |location=College Station |isbn=978-1-58544-555-4 |page=197 |edition=1st}}</ref><ref name="ALBC2009">{{cite web |title=Jacob Sheep |author=American Livestock Breeds Conservancy |author-link=American Livestock Breeds Conservancy |publisher=American Livestock Breeds Conservancy |location=Pittsboro, North Carolina |year=2009 |url=http://www.albc-usa.org/cpl/jacob.html |access-date=2011-05-05 |archive-date=2011-08-10 |archive-url=https://web.archive.org/web/20110810165333/http://www.albc-usa.org/cpl/jacob.html |url-status=live }}</ref> The unique horn structure is the only unambiguous morphological feature of bovids that distinguishes them from other [[pecorans]].<ref>{{cite journal |last=Bibi |first=F. |author2=Bukhsianidze, M. |author3=Gentry, A. |author4=Geraads, D. |author5=Kostopoulos, D. |author6=Vrba, E. |year=2009 |title=The fossil record and evolution of Bovidae: state of the field |journal=Palaeontologia Electronica |url=http://palaeo-electronica.org/2009_3/169/index.html |volume=12 |issue=3 |pages=10A |access-date=2010-11-14 |archive-date=2011-06-16 |archive-url=https://web.archive.org/web/20110616141150/http://palaeo-electronica.org/2009_3/169/index.html |url-status=live }}</ref><ref>{{cite journal |last=Gatesy |first=J. |author2=Yelon, D. |author3=DeSalle, R. |author4=Vrba, E. |year=1992 |title=Phylogeny of the Bovidae (Artiodactyla, Mammalia), based on mitochondrial ribosomal DNA sequences |journal=[[Molecular Biology and Evolution]] |volume=9 |issue=3 |pages=433–446 |pmid=1584013|doi=10.1093/oxfordjournals.molbev.a040734 |doi-access=free }}</ref> A high correlation exists between horn morphology and fighting behaviour of the individual. For instance, long horns are intended for wrestling and fencing, whereas curved horns are used in ramming.<ref name="horn1">{{cite journal |last1=Lundrigan |first1=B. |title=Morphology of horns and fighting behavior in the family Bovidae |journal=Journal of Mammalogy |volume=77 |issue=2 |pages=462–75 |doi=10.2307/1382822|jstor=1382822 |year=1996 |doi-access=free }}</ref> Males with horns directed inwards are [[Monogamy in animals|monogamous]] and solitary, while those with horns directed outwards tend to be [[Polygyny in animals|polygynous]]. These results were independent of body size.<ref name="horns2">{{cite journal |last1=Caro |first1=T. M. |last2=Graham |first2=C. M. |last3=Stoner |first3=C. J. |last4=Flores |first4=M. M. |title=Correlates of horn and antler shape in bovids and cervids |journal=Behavioral Ecology and Sociobiology |date=2003 |volume=55 |issue=1 |pages=32–41 |doi=10.1007/s00265-003-0672-6|bibcode=2003BEcoS..55...32C |s2cid=12030618 }}</ref>
Male horn development has been linked to [[Sexual selection in mammals|sexual selection]],<ref name="BroJ">{{cite journal |last=Bro-Jørgensen |first=J. |year=2007 |title=The intensity of sexual selection predicts weapon size in male bovids |journal=[[Evolution (journal)|Evolution]] |volume=61 |issue=6 |pages=1316–1326 |pmid=17542842 |doi=10.1111/j.1558-5646.2007.00111.x|s2cid=24278541 |doi-access= }}</ref><ref>{{Cite Q|Q55899859|author=Ezenwa, V.|author2=Jolles, A|author-link=Vanessa Ezenwa}}</ref> Horns are small spikes in the monogamous duikers and other small antelopes, whereas in the polygynous, they are large and elaborately formed (for example in a spiral structure, as in the [[giant eland]]). Thus, to some extent, horns depict the degree of [[Dominance (ethology)|competition]] among males in a species.<ref name="estes" /> However, the presence of horns in females is likely due to [[natural selection]].<ref name="BroJ" /><ref name="Stankowich">{{cite journal |last=Stankowich |first=T. |author2=Caro, T. |year=2009 |title=Evolution of weaponry in female bovids |journal=[[Proceedings of the Royal Society B: Biological Sciences]] |volume=276 |issue=1677 |pages=4329–34 |pmc=2817105 |pmid=19759035 |doi=10.1098/rspb.2009.1256}}</ref> The horns of females are usually smaller than those of males, and are sometimes of a different shape. The horns of female bovids are believed to have evolved for defence against predators or to express territoriality, as nonterritorial females, which are able to use [[crypsis]] for predator defence, often do not have horns.<ref name="Stankowich" /> Females possess horns only in half of the bovid genera, and females in these genera are heavier than those in the rest. Females use horns mainly for stabbing.<ref name="packer">{{cite journal |last1=Packer |first1=C. |title=Sexual Dimorphism: The Horns of African Antelopes |journal=Science |date=1983 |volume=221 |issue=4616 |pages=1191–3 |doi=10.1126/science.221.4616.1191 |pmid=17811523|bibcode=1983Sci...221.1191P |s2cid=37000507 }}</ref>
===Anatomy===
{{See also|Foregut fermentation}}
[[File:American Bison skeleton.jpg|thumb|American bison skeleton ([[Museum of Osteology]])]]
In bovids, the third and fourth [[metapodial]]s are combined into the [[cannon bone]]. The [[ulna]] and [[fibula]] are reduced, and fused with the radius and tibia, respectively. Long [[scapula]]e are present, whereas the clavicles are absent. Being [[ruminants]], the stomach is composed of four chambers: the [[rumen]] (80%), the [[omasum]], the [[Reticulum (anatomy)|reticulum]], and the [[abomasum]]. The [[ciliate]]s and [[bacteria]] of the rumen [[Fermentation|ferment]] the complex [[cellulose]] into simpler [[fatty acid]]s, which are then absorbed through the rumen wall. Bovids have a long [[small intestine]]; the length of the small intestine in [[cattle]] is {{convert|29|-|49|m|ft|abbr=on}}. Body temperature fluctuates through the day; for instance, in goats the temperature can change slightly from nearly {{convert|37|C|F}} in the early morning to {{convert|40|C|F}} in the afternoon. Temperature is regulated through sweating in cattle, whereas [[goat]]s use panting for the same. The right [[lung]], consisting of four to five [[Lobe (anatomy)|lobes]], is around 1.5 times larger than the left, which has three lobes.<ref name="adw" /><ref name="walton" />
===Dentition===
[[File:Dental pad 001.jpg|thumbnail|Dental pad of a domestic bovid: Note the absence of upper incisors and canines and the outward projection of the lower teeth.]]
Most bovids bear 30 to 32 teeth.<ref name="estes" /> While the upper [[incisor]]s are absent, the upper [[Canine tooth|canines]] are either reduced or absent. Instead of the upper incisors, bovids have a thick and tough layer of tissue, called the [[dental pad]], that provides a surface to grip grasses and foliage. They are [[hypsodont]] and [[selenodont]], since the [[molar (tooth)|molars]] and [[premolar]]s are low-[[Crown (dentistry)|crown]]ed and crescent-shaped [[Cusp (anatomy)|cusps]]. The lower incisors and canines project forward. The incisors are followed by a long toothless gap, known as the [[Diastema (dentistry)|diastema]].<ref name="EoM">{{cite book |editor=Macdonald, D. |author1=Janis, C. |author2=Jarman, P. |year=1984 |title=The Encyclopedia of Mammals |publisher=Facts on File |location=New York |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/498 498–9] |isbn=978-0-87196-871-5 |url-access=registration |url=https://archive.org/details/encyclopediaofma00mals_0/page/498 }}</ref> The general [[dentition|dental formula]] for bovids is {{DentalFormula |upper=0.0.2-3.3 |lower=3.1.3.3}}. Most members of the family are [[herbivorous]], but most duikers are [[omnivorous]]. Like other ruminants, bovids have four-chambered stomachs, which allow them to digest plant material, such as [[grass]], that cannot be used by many other animals. Ruminants (and some others like [[kangaroo]]s, [[rabbit]]s, and [[termite]]s) are able to use [[micro-organisms]] living in their guts to break down cellulose by [[fermentation (biochemistry)|fermentation]].<ref name="adw" />
==Ecology and behaviour==
[[File:Blackbuck male female.jpg|thumb|[[Blackbuck]] antelopes]]
The bovids have various methods of social organisation and social behaviour, which are classified into solitary and gregarious behaviour. Further, these types may each be divided into territorial and nonterritorial behaviour.<ref name=estes/> Small bovids such as the klipspringer, [[oribi]], and [[steenbok]] are generally solitary and territorial. They hold small territories into which other members of the species are not allowed to enter. These antelopes form [[Monogamous pairing in animals|monogamous]] pairs. Many species such as the [[dik-dik]] use [[pheromone]] secretions from the [[preorbital gland]]s and sometimes dung, as well, to mark their territories.<ref name=wyatt>{{cite book|last1=Wyatt|first1=T. D.|title=Pheromones and Animal Behaviour: Communication by Smell and Taste|url=https://archive.org/details/pheromonesanimal0000wyat|url-access=registration|date=2003|publisher=Cambridge University Press|location=Cambridge|isbn=978-0-521-48526-5|page=[https://archive.org/details/pheromonesanimal0000wyat/page/97 97]}}</ref> The offspring disperse at the time of adolescence, and males must acquire territories prior to mating.<ref name=adw/> The [[bushbuck]] is the only bovid that is both solitary and not territorial. This antelope hardly displays aggression, and tends to isolate itself or form loose herds, though in a favourable habitat, several bushbuck may be found quite close to one another.<ref name=bushbuck>{{cite web|last1=Ciszek|first1=D.|title=Bushbuck|url=http://animaldiversity.ummz.umich.edu/accounts/Tragelaphus_scriptus/|website=Animal Diversity Web|publisher=University of Michigan Museum of Zoology|access-date=28 October 2014|archive-date=28 July 2013|archive-url=https://web.archive.org/web/20130728082809/http://animaldiversity.ummz.umich.edu/accounts/Tragelaphus_scriptus/|url-status=live}}</ref>
Excluding the cephalophines (duikers), tragelaphines (spiral-horned antelopes) and the neotragines, most African bovids are gregarious and territorial. Males are forced to disperse on attaining sexual maturity, and must form their own territories, while females are not required to do so. Males that do not hold territories form bachelor herds. Competition takes place among males to acquire dominance, and fights tend to be more rigorous in limited [[rutting season]]s. With the exception of migratory males, males generally hold the same territory throughout their lives.<ref name=estes/> In the waterbuck, some male individuals, known as "satellite males", may be allowed into the territories of other males and have to wait till the owner grows old so they may acquire his territory.<ref name=waterbuck>{{cite web |last1=T. L. |first1=Newell |title=Waterbuck |url=http://animaldiversity.ummz.umich.edu/accounts/Kobus_ellipsiprymnus/ |website=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |access-date=28 October 2014 |archive-date=28 September 2012 |archive-url=https://web.archive.org/web/20120928024341/http://animaldiversity.ummz.umich.edu/accounts/Kobus_ellipsiprymnus/ |url-status=live }}</ref> [[Lek mating]], where males gather together and competitively display to potential mates, is known to exist among [[topi]]s, [[kob]]s, and [[lechwe]]s.<ref>{{cite book|author=Lott, Dale F.|year=1991|title=Intraspecific Variation in the Social Systems of Wild Vertebrates|publisher=Cambridge University Press|pages=[https://archive.org/details/intraspecificvar0000lott/page/37 37]|isbn=978-0-521-37024-0|url=https://archive.org/details/intraspecificvar0000lott/page/37}}</ref> The tragelaphines, cattle, sheep, and goats are gregarious and not territorial. In these species, males must gain absolute dominance over all other males, and fights are not confined to territories. Males, therefore, spend years in body growth.<ref name=estes/>
===Activity===
[[File:2012-wildebeest-fight.jpg|thumbnail|left|[[Blue wildebeest]] fighting for dominance]]
Most bovids are diurnal, although a few such as the buffalo, bushbuck, reedbuck, and grysbok are exceptions. Social activity and feeding usually peak during dawn and dusk. The bovids usually rest before dawn, during midday, and after dark. Grooming is usually by licking with the tongue. Rarely do antelopes roll in mud or dust. Wildebeest and buffalo usually wallow in mud, whereas the hartebeest and topi rub their heads and horns in mud and then smear it over their bodies. Bovids use different forms of vocal, olfactory, and tangible communication. These involve varied postures of neck, head, horns, hair, legs, and ears to convey sexual excitement, emotional state, or alarm. One such expression is the [[flehmen response]]. Bovids usually stand motionless, with the head high and an intent stare, when they sense danger. Some like the impala, kudu, and eland can even leap to heights of a few feet.<ref name=estes/> Bovids may roar or grunt to caution others and warn off predators.<ref name=adw/>  Bovids such as gazelles [[stotting|stot or pronk]] in response to predators, making high leaps on stiff legs, [[honest signalling|indicating honestly]] both that the predator has been seen, and that the stotting individual is strong and not worth chasing.<ref name=Bigalke1972>{{cite journal | author = Bigalke, R.C. | year = 1972 | title = Observations on the behaviour and feeding habits of the springbok ''Antidorcas marsupialis'' | journal = Zoologica Africana | volume = 7 | issue = 1 | pages = 333–359| doi = 10.1080/00445096.1972.11447448 | hdl = 10499/AJ19390 | doi-access = free }}</ref>
[[File:Springbok pronk.jpg|thumb|[[Stotting]] or pronking by a young [[springbok]] signals to [[predator]]s such as [[cheetah]]s that it is a fit and fast individual, not worth chasing.]]
In the mating season, rutting males bellow to make their presence known to females. Muskoxen roar during male-male fights, and male saigas force air through their noses, producing a roar to deter rival males and attract females. Mothers also use vocal communication to locate their calves if they get separated. During fights over dominance, males tend to [[Display (zoology)|display]] themselves in an erect posture with a level muzzle.<ref name=parker/><ref name=vaughan>{{cite book|last1=Czaplewski|first1=N. J.|last2=Ryan|first2=J. M.|last3=Vaughan|first3=T. A.|title=Mammalogy|date=2011|publisher=Jones and Bartlett Publishers|location=Sudbury|isbn=978-0-7637-6299-5|edition=5th}}</ref>
Fighting techniques differ amongst the bovid families and also depend on their build. While the hartebeest fight on knees, others usually fight on all fours. Gazelles of various sizes use different methods of combat. Gazelles usually box, and in serious fights may clash and fence, consisting of hard blows from short range. Ibex, goat and sheep males stand upright and clash into each other downwards. Wildebeest use powerful head butting in aggressive clashes. If horns become entangled, the opponents move in a circular manner to unlock them. Muskoxen will ram into each other at high speeds. As a rule, only two bovids of equal build and level of defence engage in a fight, which is intended to determine the superior of the two. Individuals that are evidently inferior to others would rather flee than fight; for example, immature males do not fight with the mature bulls. Generally, bovids direct their attacks on the opponent's head rather than its body. The S-shaped horns, such as those on the [[impala]], have various sections that help in ramming, holding, and stabbing. Serious fights leading to injury are rare.<ref name=estes/><ref name=parker>{{cite book |last1=Parker |first1=S.P. |title=Grzimek's Encyclopedia of Mammals (Volume 5) |date=1990 |publisher=McGraw-Hill Publishing |location=New York |isbn=978-0-07-909508-4 |pages=[https://archive.org/details/grzimeksencyclop0005unse/page/288 288–324, 338–9, 354–5, 432–3, 444–5, 460–1, 482–3] |edition=1st |url=https://archive.org/details/grzimeksencyclop0005unse/page/288 }}</ref><ref name="post">{{cite journal |last1=Post |first1=E. |last2=Forchhammer |first2=M. C |title=Climate change reduces reproductive success of an Arctic herbivore through trophic mismatch |journal=Philosophical Transactions of the Royal Society B: Biological Sciences|date=July 2008 |volume=363 |issue=1501 |pages=2367–2373 |doi=10.1098/rstb.2007.2207 |pmid=18006410 |pmc=2606787}}</ref>
===Diet===
[[File:Tatanka.jpg|thumbnail|left|Bovids are herbivores, feeding on grass, foliage, and plant products.]]
Most bovids alternately feed and ruminate throughout the day. While those that feed on concentrate feed and digest in short intervals, the roughage feeders take longer intervals. Only small species such as the duiker browse for a few hours during day or night.<ref name=estes/> Feeding habits are related to body size; while small bovids forage in dense and closed habitat, larger species feed upon high-fiber vegetation in open grasslands.
Subfamilies exhibit different feeding strategies. While Bovinae species graze extensively on fresh grass and diffused forage, Cephalophinae species (with the exception of ''[[Sylvicapra]]'') primarily consume fruits.<ref name=adw/> Reduncinae and Hippotraginae species depend on unstable food sources, but the latter are specially adapted to arid areas. Members of Caprinae, being flexible feeders, forage even in areas with low productivity. Tribes Alcelaphini, Hippotragini, and Reduncini have high proportions of [[monocot]]s in their diets. On the contrary, Tragelaphini and Neotragini (with the exception of ''[[Ourebia]]'') feed extensively on [[dicot]]s.<ref name=gagnon>{{cite journal |last1=Gagnon |first1=M. |last2=Chew |first2=A.E. |title=Dietary preferences in extant African Bovidae |journal=Journal of Mammalogy |date=May 2000 |volume=81 |issue=2 |pages=490–511 |doi=10.1644/1545-1542(2000)081<0490:DPIEAB>2.0.CO;2|doi-access=free }}</ref> No conspicuous relationship exists between body size and consumption of monocots.<ref>{{cite journal |last1=Sponheimer |first1=M.|last2=Lee-Thorp |first2=J.A.|last3=DeRuiter|first3=D.J.|last4=Smith |first4=J.M. |last5=van der Merwe|first5=N.J. |last6=Reed|first6=K.|last7=Grant |first7=C.C. |last8=Ayliffe |first8=L.K. |last9=Robinson |first9=T.F. |title= Diets of Southern African Bovidae: Stable Isotope Evidence |journal=Journal of Mammalogy |date=2003 |volume=84 |issue=2 |pages=471–9 |doi=10.1644/1545-1542(2003)084<0471:DOSABS>2.0.CO;2|doi-access=free }}</ref>
===Lifespan===
Most wild bovids live for 10 to 15 years. Larger species tend to live longer;<ref name=adw/> for instance, [[American bison]] can live up to 25 years and gaur up to 30 years. The mean lifespan of domesticated individuals is nearly ten years. For example, domesticated goats have an average lifespan of 12 years.
Usually males, mainly in polygynous species, have shorter lifespans than females. This can be attributed to several reasons: early dispersal of young males, aggressive male-male fights, vulnerability to predation (particularly when males are less agile, as in kudu), and malnutrition (being large in size, the male body has high nutritional requirements which may not be satisfied).<ref name=owensmith>{{cite journal|last1=Owen-Smith |first1=N. |title=Comparative mortality rates of male and female kudus: the costs of sexual size dimorphism |journal=Journal of Animal Ecology |date=1993 |volume=62 |issue=3 |pages=428–40 |doi=10.2307/5192|jstor=5192 |bibcode=1993JAnEc..62..428O |s2cid=87280870 }}</ref><ref name=toigo>{{cite journal |last1=Toigo |first1=C. |last2=Gaillard |first2=J.M. |title=Causes of Sex-Biased Adult Survival in Ungulates: Sexual Size Dimorphism, Mating Tactic or Environment Harshness? |journal=Oikos |date=2003 |volume=101 |issue=2 |pages=376–84 |doi=10.1034/j.1600-0706.2003.12073.x|bibcode=2003Oikos.101..376T }}</ref> [[Richard Despard Estes]] suggested that females mimic male secondary sexual characteristics like horns to protect their male offspring from dominant males. This feature seems to have been strongly selected to prevent male mortality and imbalanced [[sex ratio]]s due to attacks by aggressive males and forced dispersal of young males during [[adolescence]].<ref name=horn2>{{cite journal |last1=Estes |first1=R.D. |title=The significance of horns and other male secondary sexual characters in female bovids |journal=Applied Animal Behaviour Science |date=1991 |volume=29 |issue=1–4 |pages=403–51 |doi=10.1016/0168-1591(91)90264-X}}</ref>
[[File:Eland (Taurotragus oryx) male (32300677481).jpg|thumb|Eland occur in grasslands of Africa.]]
==Distribution==
Most of the diverse bovid species occur in Africa. The maximum concentration is in the [[savanna]]s of eastern Africa. Depending on their feeding habits, several species have [[adaptive radiation|radiated]] over large stretches of land, and hence several variations in dental and limb morphology are observed. Duikers inhabit the equatorial rainforests, [[sitatunga]], and lechwe occur near swamps, eland inhabit grasslands, [[springbok]] and oryx occur in deserts, [[Bongo (antelope)|bongo]] and [[anoa]] live in dense forests, and mountain goats and takin live at high altitudes.<ref name=estes/> A few bovid species also occur in Europe, Asia, and North America. Sheep and goats are found primarily in Eurasia, though the Barbary sheep and the ibex form part of the African fauna. The muskox is confined to the [[arctic tundra]]. Several bovid species have been domesticated by human beings. The domestication of goats and sheep began 10 thousand years ago, while cattle were domesticated about 7.5 thousand years ago.<ref name=adw/><ref name=Feldhamer/>
==Interaction with humans==
===Domesticated animals===
{{Further|Domestication}}
[[File:India.Mumbai.04.jpg|right|thumb|200px|[[Zebu]] oxen in [[Mumbai]]]]
The [[domestication]] of bovids has contributed to shifting the dependence of human beings from hunting and gathering to agriculture. The Bovidae includes three domesticated species whose use has spread around the world: cattle, sheep, and goats; all are from Eurasia. Other large bovids that have been domesticated but which have less ubiquitous distributions include the [[domestic buffalo]] (from the [[wild water buffalo]]), [[domestic yak]] (from the [[wild yak]]), [[zebu]] (from the [[Indian aurochs]]), [[gayal]] (from the [[gaur]]) and [[Bali cattle]] (from the [[banteng]]).<ref name=Feldhamer>{{cite book |last1=Feldhamer |first1=George A. |last2=Drickamer |first2=Lee C. |last3=Vessey |first3=Stephen H. |last4=Merritt |first4=Joseph F. |last5=Krajewski |first5=Carey |title=Mammalogy: Adaptation, Diversity, Ecology |date=2007 |publisher=Johns Hopkins University Press |isbn=978-0-8018-8695-9 |pages=519–522 |url=https://books.google.com/books?id=udCnKce9hfoC&pg=PA519 |access-date=2016-11-07 |archive-date=2023-03-14 |archive-url=https://web.archive.org/web/20230314124448/https://books.google.com/books?id=udCnKce9hfoC&pg=PA519 |url-status=live }}</ref>
Some antelopes have been domesticated including the [[oryx]]es, [[addax]], [[Taurotragus|elands]] and the extinct [[bubal hartebeest]]. In [[Ancient Egypt]] oryxes, addaxes and bubal hartebeests are depicted in carved walls.{{citation needed|date=July 2024}}
The earliest evidence of cattle domestication is from 8000 BC, suggesting that the process began in [[Cyprus]] and the [[Euphrates]] basin.<ref name=zeder>{{cite book |last1=Zeder |first1=M. A. |title=Documenting Domestication: New Genetic and Archaeological Paradigms |date=2006 |publisher=University of California Press |location=Berkeley, California |isbn=978-0-520-24638-6 |page=317}}</ref>
===Animal products===
[[File:Merino sheep.png|thumb|left|[[Merino wool]] is the most valued, with great fineness and softness.]]
[[Dairy products]] such as [[milk]], [[butter]], [[ghee]], [[yoghurt]], [[buttermilk]] and [[cheese]] are manufactured largely from domestic cattle, though the milk of sheep, goat, yak, and buffalo is also used in some parts of the world and for gourmet products. For example, buffalo milk is used to make [[mozzarella]] in Italy and ''[[gulab jamun]]'' dessert in India,<ref>{{cite web |last1=Phelan |first1=Benjamin |author2=Phelan, Benjamin |title=Others' Milk |url=http://www.slate.com/articles/life/food/2012/07/why_don_t_we_drink_other_animals_milk_the_dairy_of_camels_buffalo_pigs_sheep_and_goats_.html |publisher=Slate.com |access-date=10 October 2014 |date=24 July 2013 |archive-date=6 October 2014 |archive-url=https://web.archive.org/web/20141006124353/http://www.slate.com/articles/life/food/2012/07/why_don_t_we_drink_other_animals_milk_the_dairy_of_camels_buffalo_pigs_sheep_and_goats_.html |url-status=live }}</ref> while sheep milk is used to make blue [[Roquefort]] cheese in France.<ref>{{cite book |title=Gastronomie!: Food Museums and Heritage Sites of France |last1=Hughes |first1=Tom |last2=Hughes |first2=Meredith Sayles |year=2005 |publisher=Bunker Hill Publishing |isbn= 978-1-59373-029-1 |page=19}}</ref>
[[Beef]] is a food source high in [[zinc]], [[selenium]], [[phosphorus]], [[iron]], and [[B vitamin]]s.<ref name="whfoods">{{cite web |url=http://whfoods.org/genpage.php?pfriendly=1&tname=foodspice&dbid=141 |title=Beef, lean organic |publisher=WHFoods |date=18 October 2004 |access-date=1 April 2015 |archive-date=2 April 2015 |archive-url=https://web.archive.org/web/20150402172427/http://whfoods.org/genpage.php?pfriendly=1&tname=foodspice&dbid=141 |url-status=live }}</ref> Bison meat is lower in fat and cholesterol than beef, but has a higher protein content.<ref>{{cite web|url=http://www.bisoncentral.com/index.php?s=&c=67&d=99&a=1056&w=2&r=Y |title=National Bison Association |publisher=Bisoncentral.com |access-date=1 April 2015| archive-url= https://web.archive.org/web/20110120234037/http://bisoncentral.com/index.php?s=&c=67&d=99&a=1056&w=2&r=Y| archive-date= January 20, 2011| url-status= live}}</ref>
Bovid [[leather]] is tough and durable, with the additional advantage that it can be made into leathers of varying thicknesses - from soft clothing leather to hard shoe leather. While goat and cattle leather have a wide variety of use, [[sheepskin]] is suited only for clothing purposes.<ref>{{cite book|last1=Veldmeijer|first1=A. J.|last2=Harris|first2=S.|title=Why Leather?: The Material and Cultural Dimensions of Leather|date=2014|publisher=Sidestone Press|isbn=978-90-8890-261-1|pages=31–6}}</ref> [[Wool]] from [[Merino]] [[Domestic sheep|hoggets]] is the finest and most valuable. [[Merino wool]] is {{convert|3|-|5|in|cm|abbr=on}} long and very soft. Coarse wools, being durable and resistant to [[Pill (textile)|pilling]], are used for making tough garments<ref>{{cite web |title = Merino Sheep in Australia | url=http://www.merinos.com.au/history.asp| access-date = 1 April 2015 |archive-url = https://web.archive.org/web/20061105005633/http://www.merinos.com.au/history.asp |archive-date = 2006-11-05}}</ref> and carpets.
[[File:Drinking Horn - Brynjólfur Jónsson of Skarð, South Iceland - 1598.jpg|thumb|[[Drinking horn]] made by Brynjólfur Jónsson of Skarð, [[Iceland]], 1598]]
[[Bone meal]] is an important [[fertilizer]] rich in calcium, phosphorus, and [[nitrogen]], effective in removing [[soil acidity]].<ref name=kolay>{{cite book|last1=Kolay |first1=A. K. |title=Manures and fertilizers |date=2007 |publisher=Atlantic Publications |location=New Delhi |isbn=978-81-269-0810-3|page=98}}</ref> Bovid horns have been used as [[drinking vessel]]s since antiquity.<ref>{{Cite journal |last=Evison |first=Vera I. |date=1975 |title=Germanic Glass Drinking Horns |journal=Journal of Glass Studies |volume=17 |pages=74–87 |issn=0075-4250 |jstor=24188063}}</ref>
===In human culture===
Bovidae have featured in stories since at least the time of [[Aesop's fables]] from [[Ancient Greece]] around 600 BC. Fables by Aesop include ''[[The Crow and the Sheep]]'', ''[[The Frog and the Ox]]'', and ''[[The Wolf and the Lamb]]''.<ref>{{cite web|url=http://www.aesops-fables.com/|website=Aesop's Fables|title=Aesop's Fables|access-date=10 October 2014|archive-date=16 October 2014|archive-url=https://web.archive.org/web/20141016062054/http://www.aesops-fables.com/|url-status=live}}</ref> The mythological creature Chimera, depicted as a lion, with the head of a goat arising from its back, and a tail that might end with a snake's head, was one of the offspring of [[Typhon]] and [[Echidna (mythology)|Echidna]] and a sibling of such monsters as [[Cerberus]] and the [[Lernaean Hydra]].<ref name=peck>{{cite web|last1=Peck|title=Entry:Chimaera|url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0062:entry=chimaera-harpers|access-date=31 March 2015|archive-date=11 October 2022|archive-url=https://web.archive.org/web/20221011181119/https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0062:entry=chimaera-harpers|url-status=live}}</ref> The sheep, synonymous with the [[Goat (zodiac)|goat]] in Chinese mythology, is the eighth animal of the [[Chinese zodiac]], and a symbol of filial piety.<ref name=eberhard>{{cite book|last1=Eberhard, W.|title=A Dictionary of Chinese Symbols: Hidden Symbols in Chinese Life and Thought|year=1986|publisher=Routledge|location=London|isbn=978-0-415-00228-8|url-access=registration|url=https://archive.org/details/dictionaryofchin00wolf|author1-link=Wolfram Eberhard}}</ref>
==References==
{{Reflist|28em}}
==External links==
*{{Commons category-inline|Bovidae|''Bovidae''}}
*{{Wikispecies-inline|Bovidae|''Bovidae''}}
*{{Cite EB1911|wstitle=Bovidae |short=x}}
*{{Cite Collier's|wstitle=Bovidæ|year=1921 |short=x}}
{{Artiodactyla|R.2}}
{{Taxonbar|from=Q25497}}
{{Authority control}}
[[Category:Bovidae| ]]
[[Category:Mammal families]]
[[Category:Extant Burdigalian first appearances]]
[[Category:Taxa named by John Edward Gray]]
[[Category:Bovoidea]]