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Caprinae Temporal range: Late Miocene – present PreꞒ Ꞓ O S D C P T J K Pg N
Stone sheep (Ovis dalli stonei) in British Columbia, 2009
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Family:	Bovidae
Subfamily:	Caprinae J. E. Gray, 1821
Tribes
Caprini Ovibovini Pantholopini

The subfamily Caprinae, also sometimes referred to as the tribe Caprini, is part of the ruminant family Bovidae, and consists of mostly medium-sized bovids. A member of this subfamily is called a caprine.

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

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 1 m (3 ft) long for a full-grown grey goral (Nemorhaedus goral), to almost 2.5 m (8 ft 2 in) long for a musk ox, and from under 30 kg (66 lb) to more than 350 kg (770 lb). Musk oxen in captivity have reached over 650 kg (1,430 lb).

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

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. 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.

Species

Phylogeny based on Hassanin et al., 2009 and Calamari, 2021.

Family Bovidae

• Subfamily Caprinae or Tribe Caprini

Tribe or subtribe	Image	Genus	Species
Caprini or Caprina		Ammotragus (Blyth, 1840)	Barbary sheep, Ammotragus lervia
		Arabitragus Ropiquet & Hassanin, 2005	Arabian tahr, Arabitragus jayakari
		Budorcas Hodgson, 1850	takin, Budorcas taxicolor
		Capra 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
		Hemitragus (Hodgson, 1841)	Himalayan tahr, Hemitragus jemlahicus
		Nilgiritragus Ropiquet & Hassanin, 2005	Nilgiri tahr, Nilgiritragus hylocrius
		Oreamnos Rafinesque, 1817	mountain goat, Oreamnos americanus
		Ovis 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
		Pseudois Hodgson, 1846	bharal (Himalayan blue sheep), Pseudois nayaur
		Rupicapra Garsault, 1764	Pyrenean chamois, Rupicapra pyrenaica alpine chamois, Rupicapra rupicapra
		†Myotragus Bate, 1909	Myotragus balearicus
Ovibovini or Ovibovina		Capricornis Ogilby, 1837	Japanese serow, Capricornis crispus Chinese serow, Capricornis milneedwardsii red serow, Capricornis rubidus Sumatran serow, Capricornis sumatraensis Taiwan serow, Capricornis swinhoei Himalayan serow Capricornis thar
		Nemorhaedus Hamilton Smith, 1827	red goral, Nemorhaedus baileyi long-tailed goral, Naemorhedus caudatus gray goral, Nemorhaedus goral Chinese goral, Nemorhaedus griseus
		Ovibos Blainville, 1816	muskox, Ovibos moschatus
Pantholopini or Pantholopina		Pantholops Hodgson, 1834	Tibetan antelope, Pantholops hodgsonii

Fossil genera

The following extinct genera of Caprinae have been identified:

• 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 Tsaidamotherium †
    • Tsaidamotherium brevirostrum†
    • Tsaidamotherium hedini†

Unsorted

• †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

References

1. "Caprinae". IUCN. Archived from the original on 31 January 2022. Retrieved 31 January 2022.
2. Database, Mammal Diversity (2021-11-06), Mammal Diversity Database, doi:10.5281/zenodo.5651212, retrieved 2022-01-30
3. Gomez, W.; Patterson, T. A.; Swinton, J.; Berini, J. "Bovidae: antelopes, cattle, gazelles, goats, sheep, and relatives". Animal Diversity Web. University of Michigan Museum of Zoology. Retrieved 7 October 2014.
4. "Definition of CAPRINE". www.merriam-webster.com. Retrieved 2019-12-11.
5. Geist, Valerius (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York, NY: Facts on File. pp. 584–587. ISBN 0-87196-871-1.
6. Bover, Pere; Llamas, Bastien; Mitchell, Kieren J.; Thomson, Vicki A.; Alcover, Josep A.; Lalueza-Fox, Carles; et al. (July 2019). "Unraveling the phylogenetic relationships of the extinct bovid Myotragus balearicus (Bate 1909) from the Balearic Islands". Quaternary Science Reviews. 215: 185–195. doi:10.1016/j.quascirev.2019.05.005. S2CID 189965070.
7. Hassanin, Alexandre; Ropiquet, Anne; Couloux, Arnaud; Cruaud, Corinne (2009-04-01). "Evolution of the Mitochondrial Genome in Mammals Living at High Altitude: New Insights from a Study of the Tribe Caprini (Bovidae, Antilopinae)". Journal of Molecular Evolution. 68 (4): 293–310. Bibcode:2009JMolE..68..293H. doi:10.1007/s00239-009-9208-7. ISSN 1432-1432. PMID 19294454. S2CID 27622204.
8. Calamari, Zachary T. (June 2021). "Total Evidence Phylogenetic Analysis Supports New Morphological Synapomorphies for Bovidae (Mammalia, Artiodactyla)". American Museum Novitates (3970): 1–38. doi:10.1206/3970.1. hdl:2246/7267. ISSN 0003-0082. S2CID 235441087.
9. "Fossil Caprinae". tolweb.org.
10. "palaeos.org". Archived from the original on 2013-11-09. Retrieved 2010-08-11.

• Caprids
• Extant Miocene first appearances
• Taxa named by John Edward Gray
• Mammal subfamilies