The subfamily Caprinae, also sometimes referred to as the tribe Caprini, is part of the
ruminant family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Bovidae,
and consists of mostly medium-sized
bovid
The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the ...
s. A member of this subfamily is called a caprine, or, more informally, a goat-antelope (although they are not considered
antelope
The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia.
Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
s).
Within this tribe, a prominent clade includes
sheep
Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals 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 domesticated s ...
and
goats. 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
The Antilopines are even-toed ungulates belonging to the subfamily Antilopinae of the family Bovidae. The members of tribe Antilopini include the gazelles, blackbucks, springboks, gerenuks, dibatags, and Central Asian gazelles, are often referre ...
of the family Bovidae, with caprines being a type of bovid.
Characteristics
![Aoudad skeleton](https://upload.wikimedia.org/wikipedia/commons/6/68/Aoudad_skeleton.jpg)
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
In physical geography, tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. The term ''tundra'' comes through Russian (') from the Kildin Sámi word (') meaning "uplands", "treeless moun ...
; the
mountain goat (''Oreamnos americanus'') of
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
is specialised for very rugged terrain; the
urial (''Ovis orientalis'') occupies a largely infertile area from
Kashmir
Kashmir () is the 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 Panjal Range. Today, the term encompas ...
to
Iran
Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
, including much desert country. The
Armenian mouflon
The Armenian mouflon (''Ovis gmelini 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 Edward Bl ...
(''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
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 gree ...
, 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
The Himalayan goral (''Naemorhedus goral'') or the gray goral, is a bovid species native to the Himalayas. It is listed as Near Threatened on the IUCN Red List because the population is thought to be declining significantly due to habitat loss an ...
(''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 in size, 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
![Palaeoreas lindermayeri](https://upload.wikimedia.org/wikipedia/commons/4/49/Palaeoreas_lindermayeri.JPG)
The goat-antelope, or caprid, group is known from as early as the
Miocene
The Miocene ( ) is the first 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 means "less recen ...
, 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 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 gree ...
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 humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...
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 or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
.
Species
Phylogeny based on Hassanin ''et al.'', 2009 and Calamari, 2021.
Family
Bovidae
* Subfamily Caprinae or Tribe Caprini
Fossil genera
The following extinct genera of Caprinae have been identified:
* Tribe Caprini
** Genus ''
Myotragus''
†
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). It is one of the modern descendan ...
*** ''Myotragus balearicus''†
* Tribe Ovibovini
** Genus ''
Bootherium
''Bootherium'' (Greek: "ox" (boos), "beast" (therion)) is an extinct bovid genus from the middle to late Pleistocene of North America which contains a single species, ''Bootherium bombifrons''.McKenna & Bell, 1997, p. 442. Vernacular names for ...
''
†
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). It is one of the modern descendan ...
*** ''Bootherium bombifrons''†
** Genus ''
Euceratherium
The shrub-ox (''Euceratherium collinum'') is an extinct genus and species of ovibovine caprine native to North America along with '' Bootherium'' (''Bootherium bombifrons'') and Soergel's ox (''Soergelia mayfieldi''). Descriptions
''Eucer ...
''
†
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). It is one of the modern descendan ...
*** ''Euceratherium collinum''†
** Genus ''
Makapania
''Makapania'' is an extinct genus of large caprine or ovibovine from the Pliocene and Pleistocene of southern and East Africa. It is remarkable in that its horns were positioned laterally. Its body weight is estimated to have been about 263&nbs ...
''
†
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). It is one of the modern descendan ...
*** ''Makapania broomi''†
** Genus ''
Megalovis
''Megalovis'' is an extinct genus of bovid that lived in Eurasia during the Plio-Pleistocene.
Taxonomy
''Megalovis'' is typically considered a member of the subtribe Ovibovina of the Caprini tribe within the Antilopinae subfamily, which includ ...
''
†
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). It is one of the modern descendan ...
** Genus ''
Soergelia
''Soergelia'' is a genus of extinct ovibovine caprine that was common across Europe, North America and Asia in the Pleistocene epoch.
Species
In biology, a species is the basic unit of classification and a taxonomic rank of an organi ...
''
†
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). It is one of the modern descendan ...
*** ''Soergelia mayfieldi''†
** Genus ''
Tsaidamotherium
''Tsaidamotherium'' is an extinct genus of Late Miocene ovibovinid caprine from the Tibetan Plateau of Northwestern China. Both known species are extremely unusual in that the horns are of unequal sizes: the left horn core is several times smal ...
''
†
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). It is one of the modern descendan ...
*** ''Tsaidamotherium brevirostrum''†
*** ''Tsaidamotherium hedini''†
;Unsorted
*†''
Benicerus''
*†''
Boopsis''
*†''
Capraoryx''
*†''
Caprotragoides''
*†''
Criotherium''
*†''
Damalavus''
*†''
Gallogoral''
*†''
Lyrocerus''
*†''
Mesembriacerus''
*†''
Neotragocerus''
*†''
Nesogoral''
*†''
Norbertia''
*†''
Numidocapra''
*†''
Oioceros
''Oioceros'' is an extinct genus of spiral-horned antelope from the late Miocene. Its fossils have been found in Greece, China, Iran, and Africa. It was first discovered by Wagner in 1857, and contains nine species, ''O. rothii'', ''O. atropatene ...
''
*†''
Olonbulukia''
*†''
Pachygazella''
*†''
Pachytragus''
*†''
Palaeoreas''
*†''
Palaeoryx''
*†''
Paraprotoryx''
*†''
Parapseudotragus''
*†''
Parurmiatherium''
*†''
Praeovibos''
*†''
Procamptoceras''
*†''
Prosinotragus''
*†''
Protoryx
''Protoryx'' is an extinct genus of goat-antelope. A new species, ''P. tuvaensis'', was described by E. L. Dmitrieva and N. V. Serdyuk in 2011, from Russia
Russia (, , ), or the Russian Federation, is a List of transcontinental countrie ...
''
*† ''
Protovis
''Protovis himalayensis'' is a prehistoric species of sheep found in latest Miocene to Late Pliocene-aged strata of Himalayan Tibet in 2016.
''Protovis'' was found only by some fragments such as its horns Horns or The Horns may refer to:
* Plu ...
''
*†''
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