{{Short description|Subfamily of mammals}} {{Hatnote|Not to be confused with Antilocapridae. "Goat-antelope" redirects here. For the species sometimes called goat antelope, see Tibetan antelope.}} {{More citations needed|date=October 2015}} {{Automatic taxobox | fossil_range = {{fossil range|Late Miocene | present}} | image = Stone Sheep British Columbia.jpg | image_caption = Stone sheep (''Ovis dalli stonei'') in British Columbia, 2009 | taxon = Caprinae | authority = J. E. Gray, 1821 | subdivision_ranks = Tribes | subdivision = * Caprini * Ovibovini * Pantholopini }}
The subfamily '''Caprinae''',<ref>{{Cite web |title=Caprinae |author= |work=IUCN |date= |access-date=31 January 2022 |url=https://www.iucn.org/commissions/ssc-groups/mammals/mammals-a-e/caprinae |archive-date=31 January 2022 |archive-url=https://web.archive.org/web/20220131102816/https://www.iucn.org/commissions/ssc-groups/mammals/mammals-a-e/caprinae |url-status=dead }}</ref> also sometimes referred to as the tribe '''Caprini''',<ref>{{Citation|last=Database|first=Mammal Diversity|title=Mammal Diversity Database|date=2021-11-06|doi=10.5281/zenodo.5651212 |url=https://zenodo.org/record/5651212|access-date=2022-01-30}}</ref> is part of the ruminant family Bovidae,<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=https://animaldiversity.ummz.umich.edu/accounts/Bovidae/ |website=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |access-date=7 October 2014}}</ref> and consists of mostly medium-sized bovids. A member of this subfamily is called a '''caprine'''.<ref>{{Cite web|url=https://www.merriam-webster.com/dictionary/caprine|title=Definition of CAPRINE|website=www.merriam-webster.com|language=en|access-date=2019-12-11}}</ref>
Prominent members include sheep and goats, with some other members referred to as '''goat antelopes'''. Some earlier taxonomies considered Caprinae a separate family called Capridae (with the members being caprids), but now it is usually considered either a subfamily within the Bovidae, or a tribe within the subfamily Antilopinae of the family Bovidae, with caprines being a type of bovid.
==Characteristics== [[File:Aoudad skeleton.jpg|thumb|Skeleton of a Barbary sheep (''Ammotragus lervia'') on display at the Museum of Osteology]]
Although most goat-antelopes are gregarious and have fairly stocky builds, they diverge in many other ways – the muskox (''Ovibos moschatus'') is adapted to the extreme cold of the tundra; the mountain goat (''Oreamnos americanus'') of North America is specialised for very rugged terrain; the urial (''Ovis orientalis'') occupies a largely infertile area from Kashmir to Iran, including much desert country. The Armenian mouflon (''Ovis gmelini gmelini'') is thought to be the ancestor of the modern domestic sheep (''Ovis aries'').
Many species have become extinct since the last ice age, probably largely because of human interaction. Of the survivors: * Five are classified as endangered, * Eight as vulnerable, * Seven as of concern and needing conservation measures, but at lower risk, and * Seven species are secure.
Members of the group vary considerably in size, from just over {{convert|1|m|ft|sigfig=1|abbr=on}} long for a full-grown grey goral (''Naemorhedus goral''), to almost {{convert|2.5|m|sigfig=1|abbr=on}} long for a musk ox, and from under {{convert|30|kg|abbr=on}} to more than {{convert|350|kg|abbr=on}}. Musk oxen in captivity have reached over {{convert|650|kg|abbr=on}}.{{citation needed|date=October 2020}}
The lifestyles of caprids fall into two broad classes: 'resource-defenders', which are territorial and defend a small, food-rich area against other members of the same species; and 'grazers', which gather together into herds and roam freely over a larger, usually relatively infertile area.
The resource-defenders are the more primitive group: they tend to be smaller, dark in colour, males and females fairly alike, have long, tessellated ears, long manes, and dagger-shaped horns. The grazers (sometimes collectively known as '''tsoan caprids''', from the Hebrew ''tso'n'' meaning sheep and goats) evolved more recently. They tend to be larger, highly social, and rather than mark territory with scent glands, they have highly evolved dominance behaviours. No sharp line divides the groups, but a continuum varies from the serows at one end of the spectrum to sheep, true goats, and musk oxen at the other.
==Evolution== thumb|200px|right|''Palaeoreas lindermayeri'' fossil The goat-antelope, or caprid, group is known from as early as the Miocene, when members of the group resembled the modern serow in their general body form.<ref name=EoM>{{cite book |editor= Macdonald, D. |last=Geist |first=Valerius |year=1984 |title=The Encyclopedia of Mammals |publisher= Facts on File |location= New York, NY |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/584 584–587]|isbn= 0-87196-871-1 |url-access= registration |url=https://archive.org/details/encyclopediaofma00mals_0/page/584 }}</ref> The group did not reach its greatest diversity until the recent ice ages, when many of its members became specialised for marginal, often extreme, environments: mountains, deserts, and the subarctic region.
The ancestors of the modern sheep and goats (both rather vague and ill-defined terms) are thought to have moved into mountainous regions – sheep becoming specialised occupants of the foothills and nearby plains, and relying on flight and flocking for defence against predators, and goats adapting to very steep terrain where predators are at a disadvantage.
Internal relationships of Caprinae based on mitochondrial DNA.<ref>{{cite journal |last1=Bover |first1=Pere |last2=Llamas |first2=Bastien |last3=Mitchell |first3=Kieren J. |last4=Thomson |first4=Vicki A. |last5=Alcover |first5=Josep A. |last6=Lalueza-Fox |first6=Carles |last7=Cooper |first7=Alan |last8=Pons |first8=Joan |display-authors=6 |date=July 2019 |title=Unraveling the phylogenetic relationships of the extinct bovid ''Myotragus balearicus'' {{small|(Bate 1909)}} from the Balearic Islands |journal=Quaternary Science Reviews |lang=en |volume=215 |pages=185–195 |doi=10.1016/j.quascirev.2019.05.005 |bibcode=2019QSRv..215..185B |s2cid=189965070 |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379118306929 |url-access=subscription }}</ref>
{{clade |label1= Bovidae |1={{clade |1={{clade |label1= |1= ''Bos'' }} |label2= '''Caprinae''' |2={{clade |label1= |1={{clade |label1= |1=''Pantholops'' (Tibetan antelope) }} |2={{clade |label1= |1={{clade |label1= |1={{clade |label1= |1={{extinct}}''Bootherium'' (helmeted muskox) |2=''Ovibos'' (musk ox) }} |2={{clade |label1= |1=''Capricornis'' (serow) |2=''Naemorhedus'' (goral) }} }} |2={{clade |label1= |1={{clade |label1= |1=''Ovis'' (sheep) }} |2={{clade |label1= |1={{clade |label1= |1={{clade |label1= |1={{clade |label1= |1=''Oreamnos'' (mountain goat) }} |2={{clade |label1= |1={{clade |label1= |1=''Budorcas'' (takin) }} |2={{clade |1={{clade |1={{extinct}}''Myotragus'' (Balearic Islands goat) }} }} }} }} |2={{clade |label1= |1={{clade |label1= |1=''Rupicapra'' (chamois) }} }} }} |2={{clade |label1= |1={{clade |label1= |1={{clade |label1= |1=''Ammotragus'' (Barbary sheep) |2=''Arabitragus'' (Arabian tahr) }} }} |2={{clade |label1= |1={{clade |label1= |1=''Pseudois'' (bharal) }} |2={{clade |label1= |1=''Hemitragus'' (Himalayan tahr) |2=''Capra'' (turs, markhor, ibexes, & goats) }} }} }} }} }} }} }} }} }}
==Species== {{see also|List of bovids}}
Phylogeny based on Hassanin ''et al.'', 2009 and Calamari, 2021.<ref>{{Cite journal|last1=Hassanin|first1=Alexandre|last2=Ropiquet|first2=Anne|last3=Couloux|first3=Arnaud|last4=Cruaud|first4=Corinne|date=2009-04-01|title=Evolution of the Mitochondrial Genome in Mammals Living at High Altitude: New Insights from a Study of the Tribe Caprini (Bovidae, Antilopinae)|url=https://doi.org/10.1007/s00239-009-9208-7|journal=Journal of Molecular Evolution|language=en|volume=68|issue=4|pages=293–310|doi=10.1007/s00239-009-9208-7|pmid=19294454 |bibcode=2009JMolE..68..293H |s2cid=27622204 |issn=1432-1432|url-access=subscription}}</ref><ref name="Calamari 2021">{{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|hdl=2246/7267|hdl-access=free}}</ref><!-- Budorcas is Caprini, Naemorhedini is paraphyletic between Caprini & Ovibovini, Pantholops belongs here as basal tribe -->
:{| class="wikitable" |- ! Tribe or subtribe ! Image ! Genus ! Species |- ! rowspan="10" style="text-align:center;" | Caprini<br />or<br />Caprina | 175px |align="center"|''Ammotragus''<br />{{small|(Blyth, 1840)}} | * Barbary sheep, ''Ammotragus lervia'' |- | 175px |align="center"| ''Arabitragus''<br />{{small|Ropiquet & Hassanin, 2005}} | * Arabian tahr, ''Arabitragus jayakari'' |- | 175px |align="center"| ''Budorcas''<br />{{small|Hodgson, 1850}} | * takin, ''Budorcas taxicolor'' |- | 175px |align="center"| ''Capra''<br />{{small|Linnaeus, 1758}} | * West Asian ibex, ''Capra aegagrus'' ** bezoar ibex, ''Capra aegagrus aegagrus'' ** Sindh ibex, ''Capra aegagrus blythi'' * west Caucasian tur, ''Capra caucasica'' * east Caucasian tur, ''Capra cylindricornis'' * markhor, ''Capra falconeri'' * domestic goat, ''Capra hircus'' * alpine ibex, ''Capra ibex'' * Nubian ibex, ''Capra nubiana'' * Iberian ibex, ''Capra pyrenaica'' ** western Spanish ibex, ''Capra pyrenaica victoriae'' ** southeastern Spanish ibex, ''Capra pyrenaica hispanica'' ** †Portuguese ibex, ''Capra pyreneaica lusitanica'' ** †Pyrenean ibex, ''Capra pyreneaica pyrenaica'' * Siberian ibex, ''Capra sibirica'' * Walia ibex, ''Capra walie'' |- | 175px |align="center"| ''Hemitragus''<br />{{small|(Hodgson, 1841)}} | * Himalayan tahr, ''Hemitragus jemlahicus'' |- | 175px |align="center"| ''Nilgiritragus''<br />{{small|Ropiquet & Hassanin, 2005}} | * Nilgiri tahr, ''Nilgiritragus hylocrius'' |- | 175px |align="center"| ''Oreamnos''<br />{{small|Rafinesque, 1817}} | * mountain goat, ''Oreamnos americanus'' |- | 175px |align="center"| ''Ovis''<br />{{small|Linnaeus, 1758}} | * argali, ''Ovis ammon'' ** Marco Polo sheep, ''Ovis ammon polii'' * domestic sheep, ''Ovis aries'' * bighorn sheep, ''Ovis canadensis'' * Dall sheep, ''Ovis dalli'' * mouflon, ''Ovis gmelini'' * snow sheep, ''Ovis nivicola'' * urial, ''Ovis vignei'' |- | 175px |align="center"| ''Pseudois''<br />{{small|Hodgson, 1846}} | * bharal (Himalayan blue sheep), ''Pseudois nayaur'' |- | 175px |align="center"| ''Rupicapra''<br />{{small|Garsault, 1764}} | * Pyrenean chamois, ''Rupicapra pyrenaica'' * alpine chamois, ''Rupicapra rupicapra'' |- !rowspan="3" style="text-align:center;"| Ovibovini<br />or<br />Ovibovina | 175px |align="center"| ''Capricornis''<br />{{small|Ogilby, 1837}} | * Japanese serow, ''Capricornis crispus'' * mainland serow, ''Capricornis sumatraensis''{{efn|Calamari (2021) elevates the subspecies of mainland serow to full species status. Given IUCN classification and current Wikipedia status, they are listed as subspecies here.<ref name="Calamari 2021"/><ref>{{cite iucn |author=Phan, T.D. |author2=Nijhawan, S. |author3=Li, S. |author4=Xiao, L. |year=2020 |title=''Capricornis sumatraensis'' |volume=2020 |article-number=e.T162916735A162916910 |doi=10.2305/IUCN.UK.2020-2.RLTS.T162916735A162916910.en |access-date=28 January 2026}}</ref>}} ** Sumatran serow, ''C. s. sumatraensis'' ** Chinese serow, ''C. s. milneedwardsii'' ** Himalayan serow, ''C. s. thar'' * red serow, ''Capricornis rubidus'' * Taiwan serow, ''Capricornis swinhoei'' |- | 175px |align="center"| ''Naemorhedus''<br />{{small|Hamilton Smith, 1827}} | * red goral, ''Naemorhedus baileyi'' * long-tailed goral, ''Naemorhedus caudatus'' * gray goral, ''Naemorhedus goral'' * Chinese goral, ''Naemorhedus griseus'' |- | 175px |align="center"| ''Ovibos''<br />{{small|Blainville, 1816}} | * muskox, ''Ovibos moschatus'' |- !rowspan="1" style="text-align:center;"| Pantholopini<br />or<br />Pantholopina | 175px |align="center"| ''Pantholops''<br />{{small|Hodgson, 1834}} | * Tibetan antelope, ''Pantholops hodgsonii'' |}
==Fossil genera== The following extinct genera of Caprinae have been identified:<ref>{{Cite web|url=http://tolweb.org/Fossil_Caprinae/52476|title=Fossil Caprinae|website=tolweb.org}}</ref><ref>{{Cite web |url=http://www.palaeos.org/Caprinae |title=palaeos.org |access-date=2010-08-11 |archive-url=https://web.archive.org/web/20131109212207/http://www.palaeos.org/Caprinae |archive-date=2013-11-09 |url-status=dead }}</ref> * Tribe Caprini ** Genus ''Myotragus'' † *** ''Myotragus balearicus''† * Tribe Ovibovini ** Genus ''Bootherium'' † *** ''Bootherium bombifrons''† ** Genus ''Euceratherium'' † *** ''Euceratherium collinum''† ** Genus ''Makapania'' † *** ''Makapania broomi''† ** Genus ''Megalovis'' † ** Genus ''Soergelia'' † *** ''Soergelia mayfieldi''† ** Genus ''Speleotherium'' † *** ''Speleotherium logani''† ** Genus ''Tsaidamotherium'' † *** ''Tsaidamotherium brevirostrum''† *** ''Tsaidamotherium hedini''†
;Unsorted {{div col|colwidth=15em}} * †''Benicerus'' * †''Boopsis'' * †''Capraoryx'' * †''Caprotragoides'' * †''Criotherium'' * †''Damalavus'' * †''Gallogoral'' * †''Lyrocerus'' * †''Mesembriacerus'' * †''Neotragocerus'' * †''Nesogoral'' * †''Norbertia'' * †''Numidocapra'' * †''Oioceros'' * †''Olonbulukia'' * †''Pachygazella'' * †''Pachytragus'' * †''Palaeoreas'' * †''Palaeoryx'' * †''Paraprotoryx'' * †''Parapseudotragus'' * †''Parurmiatherium'' * †''Praeovibos'' * †''Procamptoceras'' * †''Prosinotragus'' * †''Protoryx'' * †''Protovis'' * †''Pseudotragus'' * †''Qurliqnoria'' * †''Samotragus'' * †''Sinocapra'' * †''Sinomegoceros'' * †''Sinopalaeoceros'' * †''Sinotragus'' * †''Sivacapra'' * †''Sporadotragus'' * †''Tethytragus'' * †''Tossunnoria'' * †''Turcocerus'' * †''Urmiatherium'' {{div col end}}
==Notes== {{notelist}}
==References== {{reflist|25em}}
{{Artiodactyla|R.3}} {{Taxonbar|from=Q189804}} {{Authority control}}
Category:Caprinae Category:Extant Miocene first appearances Category:Taxa named by John Edward Gray Category:Mammal subfamilies