The subfamily Caprinae, also sometimes referred to as the tribe Caprini, is part of the
ruminant
Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
family
Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Bovidae
The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
,
and consists of mostly medium-sized
bovid
The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
s. A member of this subfamily is called a caprine.
Prominent members include
sheep
Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
and
goat
The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
s, with some other members referred to as goat antelopes. Some earlier
taxonomies
image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy
Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
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
Antilopinae is a subfamily of even-toed ungulates in the family Bovidae. The members of tribe Antilopini are often referred to as true antelopes, and include the gazelles, blackbucks, springboks, gerenuks, dibatags, and Central Asian gazelles. ...
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
The muskox (''Ovibos moschatus'') is a hoofed mammal of the family Bovidae. Native to the Arctic, it is noted for its thick coat and for the strong odor emitted by males during the seasonal rut, from which its name derives. This musky odor ha ...
(''Ovibos moschatus'') is adapted to the extreme cold of the
tundra
In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...
; the
mountain goat
The mountain goat (''Oreamnos americanus''), also known as the Rocky Mountain goat, is a cloven-footed mammal that is endemic to the remote and rugged mountainous areas of western North America. A subalpine to truly alpine species, it is a s ...
(''Oreamnos americanus'') of
North America
North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
is specialised for very rugged terrain; the
urial
The urial ( ; ''Ovis vignei''), also known as arkars, shapo, or shapu, is a wild sheep native to Central and South Asia. It is listed as Vulnerable on the IUCN Red List.
Taxonomy
''Ovis vignei'' was the scientific name proposed by Edward Bly ...
(''Ovis orientalis'') occupies a largely infertile area from
Kashmir
Kashmir ( or ) is the Northwestern Indian subcontinent, northernmost geographical region of the Indian subcontinent. Until the mid-19th century, the term ''Kashmir'' denoted only the Kashmir Valley between the Great Himalayas and the Pir P ...
to
Iran
Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
, including much desert country. The
Armenian mouflon
The Armenian mouflon (''Ovis orientalis gmelini'') is an endangered subspecies of mouflon endemic to Iran, Armenia, Nakhchivan Autonomous Republic of Azerbaijan, Turkey and Iraq.
Taxonomy
The Armenian mouflon was first described in 1840 by Edwar ...
(''Ovis gmelini gmelini'') is thought to be the ancestor of the modern
domestic sheep
Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to ...
(''Ovis aries'').
Many species have become extinct since the last
ice age
An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
, 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 long for a full-grown
grey goral (''Nemorhaedus goral''), to almost long for a musk ox, and from under to more than . Musk oxen in captivity have reached over .
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
The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, when members of the group resembled the modern
serow
The serow (, or ), is any of four species of medium-sized goat-like or antelope-like mammals in the genus ''Capricornis''. All four species of serow were, until recently, classified under ''Naemorhedus'', which now only contains the gorals.
Exta ...
in their general body form.
The group did not reach its greatest diversity until the recent
ice age
An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
s, when many of its members became specialised for marginal, often extreme, environments: mountains, deserts, and the
subarctic
The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of hemiboreal regions and covering much of Alaska, Canada, Iceland, the north of Fennoscandia, Northwestern Russia, Siberia, and the Cair ...
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
Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
.
Species
Phylogeny based on Hassanin ''et al.'', 2009 and Calamari, 2021.
Family
Bovidae
The Bovidae comprise the family (biology), biological family of cloven-hoofed, ruminant mammals that includes Bos, cattle, bison, Bubalina, buffalo, antelopes (including Caprinae, goat-antelopes), Ovis, sheep and Capra (genus), goats. A member o ...
* Subfamily Caprinae or Tribe Caprini
:
Fossil genera
The following extinct genera of Caprinae have been identified:
* Tribe Caprini
** Genus ''
Myotragus
''Myotragus'' (Neo-Latin, derived from the Greek: , "mouse-goat") is an extinct genus of goat-antelope in the tribe Caprini which lived on the Balearic Islands of Mallorca and Menorca in the western Mediterranean until its extinction around 4 ...
''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Myotragus balearicus''†
* Tribe Ovibovini
** Genus ''
Bootherium
''Bootherium'' (Greek: "ox" (boos), "beast" (therion)) is an extinction, extinct bovid genus from the Middle Pleistocene, Middle to Late Pleistocene of North America which contains a single species, ''Bootherium bombifrons''.McKenna & Bell, 1997, ...
''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Bootherium bombifrons''†
** Genus ''
Euceratherium''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Euceratherium collinum''†
** Genus ''
Makapania''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Makapania broomi''†
** Genus ''
Megalovis''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
** Genus ''
Soergelia''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Soergelia mayfieldi''†
** Genus ''
Tsaidamotherium''
†
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
*** ''Tsaidamotherium brevirostrum''†
*** ''Tsaidamotherium hedini''†
;Unsorted
* †''
Benicerus''
* †''
Boopsis''
* †''
Capraoryx''
* †''
Caprotragoides''
* †''
Criotherium''
* †''
Damalavus''
* †''
Gallogoral''
* †''
Lyrocerus
''Lyrocerus'' is an extinct genus of caprine bovid that inhabited Eurasia during the Neogene period.
Distribution
''Lyrocerus sata'' is known from China
China, officially the People's Republic of China (PRC), is a country in East ...
''
* †''
Mesembriacerus''
* †''
Neotragocerus''
* †''
Nesogoral''
* †''
Norbertia''
* †''
Numidocapra
''Numidocapra'' is an extinct genus of bovid from the Pleistocene of Africa.
Taxonomy
Originally placed in the subfamily caprinae, ''Numidocapra'' is now generally classified as a member of the subfamily Alcelaphinae. Three species are conside ...
''
* †''
Oioceros''
* †''
Olonbulukia''
* †''
Pachygazella''
* †''
Pachytragus''
* †''
Palaeoreas''
* †''
Palaeoryx''
* †''
Paraprotoryx''
* †''
Parapseudotragus''
* †''
Parurmiatherium''
* †''
Praeovibos
''Praeovibos'', also known as the giant muskox, is an extinct genus of bovid that contains a single species, ''Praeovibos priscus''. It is closely related to the living muskox (''Ovibos moschatus''), and is placed as a part of the same tribe Ovib ...
''
* †''
Procamptoceras''
* †''
Prosinotragus''
* †''
Protoryx''
* †''
Protovis''
* †''
Pseudotragus''
* †''
Qurliqnoria''
* †''
Samotragus''
* †''
Sinocapra''
* †''
Sinomegoceros''
* †''
Sinopalaeoceros''
* †''
Sinotragus''
* †''
Sivacapra''
* †''
Sporadotragus''
* †''
Tethytragus''
* †''
Tossunnoria''
* †''
Turcocerus''
* †''
Urmiatherium''
References
{{Authority control
Extant Miocene first appearances
Taxa named by John Edward Gray
Mammal subfamilies