Fact-checked by Grok 2 weeks ago

Caprinae

Caprinae is a subfamily of the ruminant family Bovidae, consisting primarily of medium-sized bovids such as sheep, goats, chamois, serows, gorals, tahrs, and muskoxen, renowned for their remarkable adaptations to steep, rocky, and high-altitude environments.^[1] This diverse group encompasses approximately 35 extant species, which are distributed across mountainous regions of Europe, Asia, Africa, and North America, with some species also inhabiting arid grasslands and forests.^[1]^[2] Members of Caprinae exhibit a range of body sizes, from the small goral weighing around 20–30 kg to the robust muskox reaching up to 400 kg, and are distinguished by their cloven hooves, four-chambered stomachs for ruminating vegetation, and often sexually dimorphic horns used for defense and display.^[3]^[4] Taxonomically, Caprinae is divided into several tribes, including Caprini (true goats and sheep), Rupicaprini (goat-antelopes like chamois and tahrs), and Ovibovini (muskox), reflecting evolutionary divergences that began in the Miocene epoch, with fossils indicating origins in Eurasia before widespread dispersal.^[4] These animals are highly agile climbers and jumpers, enabling them to navigate precipitous cliffs and escape predators, while their thick woolly coats provide insulation against extreme cold in alpine habitats.^[5] Socially, most species form herds, particularly females with young, exhibiting behaviors such as seasonal migrations between summer and winter ranges to optimize foraging on grasses, shrubs, and lichens.^[1] Conservation concerns are prominent, as many Caprinae species face threats from habitat loss, poaching, and climate change, leading to several being listed as vulnerable or endangered by the IUCN as of 2024.^[6]

Taxonomy

Classification

Caprinae is classified as a subfamily within the family Bovidae, which belongs to the order Artiodactyla and suborder Ruminantia.^[7]^[8] This placement reflects the group's position among even-toed ungulates characterized by their ruminant digestive system and even number of toes.^[7] The subfamily Caprinae was first formally described by the British zoologist John Edward Gray in 1821, establishing its binomial nomenclature within the Bovidae framework.^[9] Gray's designation laid the foundation for recognizing Caprinae as a distinct group of goat-like and sheep-like bovids, separate from other subfamilies such as Bovinae or Antilopinae.^[9] This initial taxonomy has been refined over time through morphological and genetic analyses. In contemporary classification, the tribe Caprini within Caprinae encompasses genera such as Capra (goats) and Ovis (sheep).^[10] Molecular data, including cytochrome b sequences, have largely supported the monophyly of Caprinae, though debates persist regarding the inclusion of certain taxa like Saiga, with some studies excluding it to maintain clade integrity.^[11] Seminal works, such as Gentry (1992) on bovid subfamilies and Gatesy et al. (1997) on molecular systematics, have bolstered this view by integrating morphological and genetic evidence to affirm Caprinae's cohesive evolutionary lineage.^[12]^[11]

Subdivisions

The subfamily Caprinae is divided into four main tribes: Pantholopini, Ovibovini, Rupicaprini, and Caprini, reflecting phylogenetic relationships based on morphological and genetic data.^[1] The inclusion of Pantholopini (chiru) in Caprinae is supported by molecular data but sometimes debated, with proposals for a separate subfamily (Pantholopinae). The tribe Caprini encompasses true goats (Capra), sheep (Ovis), and related forms like blue sheep (Pseudois) and aoudads (Ammotragus), representing the core goat-like and sheep-like lineages adapted to rugged terrains. In contrast, minor tribes such as Ovibovini include more specialized forms like the muskox (Ovibos) and takin (Budorcas), which exhibit bovine-like features alongside caprine traits.^[3] These divisions highlight the subfamily's diversity, with Caprini comprising the majority of species and genera.^[1] As of 2023, Caprinae includes approximately 12 living genera and around 35 extant species, with high levels of endemism in mountainous regions such as the Himalayas, where species like the Himalayan tahr (Hemitragus jemlahicus) are restricted.^[1]^[13] The following table summarizes key living genera, their tribal affiliations, and approximate species counts based on recent assessments:

Tribe	Genus	Common Name Examples	Approximate Species Count
Pantholopini	Pantholops	Tibetan antelope (chiru)	1
Ovibovini	Ovibos	Muskox	1
Ovibovini	Budorcas	Takin	1
Rupicaprini	Naemorhedus	Gorals	4
Rupicaprini	Capricornis	Serows	5
Rupicaprini	Rupicapra	Chamois	2
Caprini	Capra	Ibexes, markhor, tur	9
Caprini	Ovis	Argali, bighorn sheep, urial	6
Caprini	Ammotragus	Aoudad (Barbary sheep)	1
Caprini	Pseudois	Blue sheep, dwarf blue sheep	2
Caprini	Oreamnos	Mountain goat	1
Caprini	Arabitragus	Arabian tahr	1
Caprini	Nilgiritragus	Nilgiri tahr	1
Caprini	Hemitragus	Himalayan tahr	1

This classification accounts for 35 species in total.^[1]^[14] Taxonomic revisions in the 2000s and 2010s, driven by molecular phylogenetics, have refined these subdivisions. For instance, the genus Hemitragus was revised, with the Arabian tahr moved to Arabitragus and the Nilgiri tahr to Nilgiritragus, leaving Hemitragus monospecific for the Himalayan tahr, based on evidence of polyphyly within the group.^[15] Similarly, genetic studies in the 2010s elevated subspecies to full species status in Naemorhedus (gorals, now four species) and Capricornis (serows, now five species), addressing prior underestimation of diversity in these Asian endemics.^[14] The genus Pseudois remains with two recognized species (blue sheep and dwarf blue sheep), though ongoing genetic analyses continue to inform potential further adjustments.^[14] These changes underscore the role of DNA-based evidence in clarifying evolutionary relationships within Caprinae.^[16]

Physical Characteristics

Morphology

Caprinae species display considerable variation in body size, ranging from approximately 20 kg in smaller forms like the goral (Naemorhedus goral) to over 300 kg in larger taxa such as the muskox (Ovibos moschatus), with most falling between 20 and 300 kg.^[17] Sexual dimorphism is pronounced, particularly in body mass and horn size, where males are typically substantially larger than females, with body mass ratios of 1.5–1.8 (i.e., 50–80% heavier) in many wild species and exhibit more robust skeletal structures to support larger horns.^[17] ^[18] This dimorphism arises from evolutionary pressures related to male-male competition, though it varies across genera, with higher degrees observed in species like wild goats (Capra spp.).^[18] The overall body plan is adapted for agility in rugged terrains, featuring a cylindrical torso, relatively short tail, and sturdy limbs that facilitate climbing and leaping.^[17] A key digestive adaptation is the four-chambered stomach, consisting of the rumen, reticulum, omasum, and abomasum, which enables rumination—the process of regurgitating and rechewing food to break down fibrous plant material efficiently.^[19] Horns are a defining feature, composed of a bony core covered by a keratinous sheath that grows continuously throughout life and remains permanent, unlike antlers; they are present in both sexes in many species, though males often have longer, thicker horns for defense and display.^[20] The skull exhibits a characteristic convex profile with prominent supraorbital ridges and large orbits that provide wide fields of vision, essential for detecting predators in open or mountainous habitats.^[21] The dental formula is I 0/4, C 0/0, P 3/3, M 3/3, reflecting the absence of upper incisors and canines, with the lower canines modified to function as incisors alongside three true lower incisors for grazing.^[22] Coat morphology varies by genus: sheep (Ovis spp.) typically have dense, woolly underfur for insulation, while goats (Capra spp.) possess coarser guard hairs; both undergo seasonal molting, shedding thicker winter coats in spring to reveal sleeker summer pelage.^[23] This molting is driven by photoperiod changes and helps regulate thermoregulation across diverse climates.^[24]

Adaptations

Caprinae species exhibit remarkable morphological and physiological adaptations that facilitate survival in rugged, high-altitude, and often harsh environments. These traits enable them to navigate steep terrains, maintain homeostasis under extreme temperatures, detect threats efficiently, and extract nutrients from fibrous vegetation. For agile climbing on rocky terrains, Caprinae possess specialized hooves with a hard outer edge for chipping into surfaces and a soft, rubbery inner pad that provides traction and absorbs shock on uneven rocks.^[25] This structure, combined with highly flexible joints in the legs, allows species like the Alpine ibex (Capra ibex) to scale near-vertical cliffs with precision and stability.^[26] Additionally, their countershaded pelage—darker on the upper body and lighter below—enhances camouflage against rocky backgrounds, reducing visibility to predators in sparse, mountainous vegetation. Thermoregulation in Caprinae is supported by dense underwool in high-altitude species, such as ibex, which forms an insulating layer of fine, interwoven fibers that traps air and retains body heat during cold exposure.^[27] In winter, many Caprinae, including the Alpine ibex, reduce their metabolic rates by more than 50% through hypometabolism, minimizing endogenous heat production and conserving energy in low-resource, cold climates.^[25] Sensory enhancements are crucial for predator detection in open, exposed habitats. Caprinae have panoramic vision with a field of approximately 320–340 degrees, enabled by horizontally positioned rectangular pupils that maximize peripheral awareness while grazing or scanning for threats.^[28] Their acute hearing, sensitive to a wide frequency range beyond human capabilities (up to 40 kHz in goats), allows early detection of approaching predators through low-frequency footfalls or vocalizations.^[29] Digestive efficiencies in Caprinae rely on foregut fermentation in the rumen, where symbiotic microbes break down high-fiber plant material into volatile fatty acids, enabling sustenance on coarse, low-quality forage typical of alpine meadows.^[30] While the rumen handles the majority of fiber digestion, the caecum provides supplementary fermentation in some genera, contributing to additional nutrient extraction from undigested residues.^[31]

Distribution and Habitat

Geographic Range

The subfamily Caprinae is native to the Holarctic biogeographic region, encompassing much of the northern hemisphere's temperate and mountainous zones.^[1] Their distribution is dominated by the Palearctic realm, which includes Eurasia from the Iberian Peninsula—home to species like the Iberian ibex (Capra pyrenaica)—extending eastward through diverse terrains to the Himalayas, where taxa such as the blue sheep (Pseudois nayaur) and Himalayan tahr (Hemitragus jemlahicus) occur.^[1]^[2] This realm also incorporates North Africa, supporting species like the Barbary sheep (Ammotragus lervia) in rocky habitats across the Atlas Mountains and Sahara fringes, as well as the Walia ibex (Capra walie) in the Ethiopian highlands.^[1] In the Nearctic realm, Caprinae presence is limited but significant, with North American endemics such as the bighorn sheep (Ovis canadensis) and mountain goat (Oreamnos americanus) occupying the Rocky Mountains and coastal ranges, reflecting historical faunal exchanges via the Bering land bridge during Pleistocene lowstands.^[1]^[32] Introduced populations have expanded Caprinae beyond their native ranges, primarily through human-mediated translocations of domestic goats (Capra hircus) that later feralized. In Australia, goats arrived with the First Fleet in 1788 and established widespread feral herds across arid and semi-arid regions by the 19th century.^[33] Similarly, introductions to New Zealand began in the 1770s by European explorers, leading to feral populations in forested and alpine areas.^[34] In Hawaii, goats were brought by Captain James Cook in 1778, rapidly proliferating on volcanic islands and causing ecological disruptions.^[35] These non-native populations now number in the millions in some areas, though management efforts continue to mitigate their impacts. Human activities have increasingly fragmented Caprinae ranges, particularly in the 20th century, through habitat conversion, overhunting, and infrastructure development. In the Mediterranean Basin, species like the Iberian ibex experienced severe declines, with populations reduced to very low numbers in some regions, such as fewer than 20 individuals by the early 20th century, due to agricultural expansion and poaching, leading to localized extirpations and isolated remnants.^[36] The Pyrenean subspecies (Capra pyrenaica pyrenaica), for instance, became extinct in 2000 following century-long fragmentation exacerbated by human pressures.^[37] Such patterns underscore broader vulnerabilities in densely populated Eurasian and North African zones, where connectivity between subpopulations has diminished.

Preferred Environments

Caprinae species predominantly favor rugged terrains such as alpine meadows, sheer cliffs, and montane forests, often at elevations exceeding 2,000 meters, where steep slopes and rocky outcrops provide escape routes from predators and access to forage.^[38] These environments allow for agile navigation, with species like the Alpine ibex (Capra ibex) utilizing precipitous rocky areas above the treeline for security and grazing on sparse vegetation. Mountain goats (Oreamnos americanus) similarly select cliff-dominated habitats in alpine zones for their sure-footedness on near-vertical surfaces.^[39] Climate tolerances among Caprinae span temperate to subarctic conditions, enabling occupancy of diverse regions from the European Alps to Arctic tundra fringes, though most thrive in cool, montane climates with seasonal snow cover.^[1] Notable exceptions include adaptations to arid and desert environments, as seen in the Barbary sheep (Ammotragus lervia), which inhabits semi-arid mountainous areas of North Africa, enduring hot, dry conditions in rocky steppes and canyons with minimal water availability. Many Caprinae engage in seasonal microhabitat shifts, migrating altitudinally between lowlands and highlands to optimize foraging opportunities, descending to lower elevations in winter for reduced snow depth and ascending to summer alpine meadows for fresh herbaceous growth.^[40] For instance, bighorn sheep (Ovis canadensis) in North American ranges move upslope during warmer months to exploit nutrient-rich grasses, returning to valley bottoms when high-elevation forage diminishes.^[41] Caprinae typically graze in open grasslands and scrublands maintained through herbivore activity, which promotes vegetative regrowth and diversity, while generally avoiding dense forests that limit mobility and visibility.^[4] This preference fosters symbiotic dynamics in grassland ecosystems, where their browsing and grazing help sustain open habitats suitable for other herbivores, as observed in mixed-species assemblages in montane steppes.^[42]

Behavior and Ecology

Caprinae species typically form social groups that vary by sex, age, and season, with females and their offspring comprising the core of stable, matriarchal herds ranging from 5 to 20 individuals. These female-led groups provide protection and facilitate cooperative foraging in rugged terrains, while adult males outside the breeding season often remain solitary or join loose bachelor herds of 2 to 10 individuals to minimize conflict and conserve energy. During the rut, males may integrate into female groups temporarily, but sexual segregation predominates year-round in many species, such as bighorn sheep (Ovis canadensis), where matrilineal bonds drive long-term female associations and philopatry.^[43]^[44] Dominance hierarchies within Caprinae groups are well-defined, particularly among males, where linear rankings are established through ritualized horn clashes and displays that assess strength without severe injury. In species like bighorn sheep, these hierarchies determine access to resources and mates, with older, larger males occupying top positions; females also maintain stable, age-correlated hierarchies, though less linear and aggressive than in males. Social dominance in goats (Capra spp.), including feral populations, correlates with horn size and is reinforced by head-butting, influencing feeding priority and group position.^[45]^[46]^[47] Communication among Caprinae relies on a multimodal system including vocalizations such as bleats and grunts for alarm or affiliation, scent marking via preorbital glands or urine to signal presence and status, and postural displays like head tosses or ear positions to convey dominance or submission. These signals maintain group cohesion and reduce aggression, as seen in sheep where visual cues like stomping precede conflicts.^[48]^[49]^[50] Territoriality in Caprinae varies markedly, with goats exhibiting milder, resource-based defense through vocal threats and brief chases rather than fixed boundaries, whereas mountain sheep like bighorn display more aggressive territorial defense during the rut, where dominant males clash to secure harems or mating access to groups of females. This contrast reflects adaptations to habitat: goats in fragmented ranges prioritize mobility, while sheep in open alpine areas enforce hierarchies to control prime foraging sites.^[5]^[51]

Diet and Reproduction

Members of the Caprinae subfamily are primarily herbivorous, exhibiting a mixed feeding strategy that includes both grazing on grasses and browsing on shrubs, forbs, and lichens, depending on seasonal availability and habitat. In summer, species like mountain goats (Oreamnos americanus) preferentially consume grasses and forbs, while winter diets shift to more fibrous materials such as shrubs and lichens to meet nutritional needs in resource-scarce environments.^[39] Selective feeding allows them to support rumen fermentation efficiency typical of ruminant herbivores. For example, desert bighorn sheep (Ovis canadensis nelsoni) incorporate a diverse array of up to 41 plant species, balancing grasses, forbs, and shrubs to optimize energy intake.^[52] Reproductive cycles in Caprinae are typically seasonally polyestrous, with most species entering estrus multiple times during a fall-to-winter breeding period influenced by photoperiod, though cycle lengths vary from 16-17 days in sheep to 21 days in goats.^[53] Breeding often peaks in a fall rut, leading to gestation periods typically of 150-180 days in most species such as sheep and goats, though longer in some like the muskox (up to 240 days), resulting in births synchronized to spring for optimal foraging conditions.^[54]^[55] Twinning is rare in wild populations, occurring periodically in most species with usually one offspring per year, though rates can reach 50-100% in select domestic breeds under managed conditions.^[42]^[56] Parental care involves immediate following behavior by precocial young, such as lambs or kids, which remain closely attached to their mothers for protection and nursing in social groups where foraging occurs collectively.^[57] Weaning typically happens at 3-6 months, allowing offspring to transition to solid forage while minimizing maternal energy costs.^[58] Seasonal diet shifts significantly influence reproductive success, as higher nutritional quality in spring—through access to fresh grasses and forbs—enhances lamb survival rates by improving maternal condition and milk production, with studies on bighorn sheep showing positive correlations between late-winter fat reserves and summer offspring viability.^[59]^[60]

Evolution and Fossil Record

Origins and Phylogeny

The subfamily Caprinae originated in Eurasia during the early Miocene, approximately 18–15 million years ago, evolving from early bovid ancestors within the broader Aegodontia group.^[1] This emergence coincided with significant climatic and physiographic changes, including the closure of the Tethys seaway, which facilitated the diversification of ruminants across the region.^[61] Fossil evidence indicates early representatives of related bovid subfamilies appearing in both African and Eurasian deposits around 14 million years ago. Phylogenetic reconstructions, drawing on both mitochondrial DNA (mtDNA) sequences like cytochrome b and nuclear genes, strongly support the monophyly of Caprinae as a distinct clade within Bovidae.^[62] Within this clade, the tribe Caprini—encompassing genera such as Ovis and Capra—forms a robustly supported subgroup, with divergence estimates placing the split between Ovis and Capra at 5–7 million years ago based on allozyme and sequence data calibrated against geological timelines. These molecular analyses highlight a pattern of gradual basal branching followed by more recent intrageneric radiations. Major evolutionary events in Caprinae history include Pleistocene radiations driven by glacial-interglacial cycles, which promoted adaptive expansions into Holarctic environments as retreating ice sheets opened new habitats.^[63] Molecular clock methods, calibrated to key fossil nodes such as middle Miocene bovid divergences, reveal accelerated speciation rates in the late Miocene to Pliocene, particularly along the northern margins of the rising Tibetan Plateau (Himalayas), where tectonic uplift created diverse montane niches for stem caprines.^[64] This rapid diversification is evidenced by divergence times around 8–13 million years ago for major caprine lineages, underscoring the role of orogenic processes in shaping the subfamily's phylogeny.^[65]

Extinct Genera

The fossil record of Caprinae documents a diverse array of extinct genera spanning from the late Miocene to the Holocene, that illuminate the subfamily's evolutionary diversification across Eurasia, North America, and beyond.^[9] These genera highlight adaptations to varied paleoenvironments, from mountainous terrains to open steppes, and provide key insights into the biogeographic spread and morphological evolution of caprines.^[9] Among the major extinct genera is Sinocapra, an early representative of the tribe Caprini known from the Pliocene of northern China and North America, with species such as S. minor from Shanxi Province, China (~3.5 Ma), and S. willdownsi from Nevada, USA (4.95–4.50 Ma), indicating migrations from Asia to North America in the earliest Pliocene. Fossils reveal goat-like features such as small, straight to slightly curved horn cores and robust metapodials suited for rugged terrain, suggesting it occupied forested or hilly habitats in subtropical Asia.^[9] Similarly, Oioceros from the late Miocene of Europe, with key specimens from Pikermi in Greece, exemplifies primitive caprine morphology through its spiral-horned structure and dental traits akin to early sheep-goats, indicating a role in the initial radiation of the subfamily across the Old World.^[66] Notable among Pleistocene forms are the ovibovines, precursors to modern muskoxen (Ovibos), such as Bootherium and Symbos from North America, which adapted to Ice Age tundra-steppe ecosystems with thick skulls for head-butting and dense woolly pelage inferred from associated remains. These genera, documented in sites like the Yukon and Alaska, reached body sizes up to 300 kg and demonstrate Holarctic dispersal patterns.^[67] Giant forms like Soergelia, an ovibovine caprine widespread in the Pleistocene of Europe, Asia, and North America, are represented by robust cranial and postcranial fossils from sites such as Monte Argentario in Italy, showcasing enlarged horn cores and limb bones indicative of grazing in open grasslands during glacial maxima.^[68] Extinction patterns within Caprinae align with broader late Pleistocene megafaunal losses, particularly around 12,000–10,000 years ago, affecting genera adapted to steppe environments amid climatic shifts and human pressures; examples include the insular dwarf Myotragus balearicus from the Balearic Islands, which persisted into the early Holocene before vanishing circa 4,000 years ago due to isolation and arrival of humans.^[69]^[70] These events reduced caprine diversity, with surviving lineages showing phylogenetic continuity to extant forms while highlighting the subfamily's vulnerability to end-Pleistocene environmental upheavals.^[71]

Conservation Status

Threats

Caprinae species face significant threats from habitat loss primarily driven by deforestation and agricultural expansion, which have encroached upon their montane and alpine habitats worldwide. Livestock grazing and conversion of grasslands to farmland further exacerbate resource competition, leading to degradation of foraging areas and increased human-wildlife conflict in regions like Central Asia and the Himalayas.^[72] Poaching and illegal hunting for meat, horns, and trophies remain a critical danger, particularly for species valued in traditional medicine and sport. The argali (Ovis ammon), for instance, has experienced substantial population declines across its range in Central Asia due to persistent poaching, despite legal protections, contributing to its threatened status.^[73] Climate change poses an emerging threat by altering alpine ecosystems through rising temperatures and shifting vegetation zones, forcing high-elevation Caprinae like ibex and chamois to adapt to reduced snow cover and changing forage availability. These shifts disrupt migration patterns and habitat suitability, amplifying vulnerability for montane specialists.^[74] Disease transmission from domestic livestock to wild Caprinae represents another key risk, with outbreaks of foot-and-mouth disease (FMD) in Asia during the 2000s severely impacting populations of wild sheep and goats through direct contact and shared grazing areas. Serotype Asia 1 epidemics, originating in southern Asia, spread rapidly among cloven-hoofed wildlife, highlighting the dangers of interspecies pathogen spillover.^[75]

Protection Efforts

Conservation efforts for Caprinae species have focused on mitigating threats through protected areas, international trade regulations, and active population management. According to the IUCN Red List, approximately 25% of the roughly 35 recognized Caprinae species are classified as threatened (Vulnerable, Endangered, or Critically Endangered), highlighting the need for targeted interventions. For instance, the markhor (Capra falconeri) is listed as Near Threatened due to population recoveries driven by conservation actions, though it remains vulnerable to poaching in parts of its range. Similarly, the Arabian tahr (Arabitragus jayakari) is categorized as Endangered, with its fragmented populations in the Arabian Peninsula numbering fewer than 2,500 mature individuals. The Walia ibex (Capra walie) is assessed as Vulnerable, confined to a small area in Ethiopia's Simien Mountains, where a 2025 survey estimated fewer than 250 mature individuals, indicating ongoing declines that meet criteria for Critically Endangered and may warrant uplisting.^[76] Establishment of protected areas has been crucial for safeguarding Caprinae habitats and populations. In Pakistan, Khunjerab National Park, created in 1975, spans over 2,200 square kilometers in the Karakoram Mountains and serves as a key refuge for the markhor and Marco Polo sheep (Ovis ammon polii), with anti-poaching patrols and habitat monitoring contributing to local population stability. Other reserves, such as those in the Himalayas and Alps, similarly protect species like the blue sheep (Pseudois nayaur) and chamois (Rupicapra rupicapra) from habitat fragmentation and human encroachment. International agreements have played a pivotal role in curbing illegal trade in Caprinae horns and trophies. The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), effective since 1975, lists numerous Caprinae taxa on Appendix I, prohibiting commercial international trade for species like the markhor and most ibex subspecies, while Appendix II regulates trade for others such as argali (Ovis ammon) to prevent overexploitation. These listings have reduced poaching pressures, particularly for medicinal and ornamental uses, leading to measurable population upticks in regulated regions. Reintroduction programs have demonstrated significant success in restoring Caprinae populations across North America. For the Rocky Mountain bighorn sheep (Ovis canadensis canadensis), extensive translocations since the 1950s have increased numbers from an estimated 20,000–25,000 continent-wide to over 80,000 as of 2025, a more than 300% rise attributed to habitat protection and disease management. Similar efforts for desert bighorn sheep in the southwestern United States have bolstered subpopulations, emphasizing the role of captive breeding and monitoring in overcoming historical declines.

References

[1]
Subfamily Caprinae - Sheep, goats, and relatives - Ultimate Ungulate
Feb 24, 2023 · The Caprinae is one of the most successful bovid subfamilies, with 35 currently recognized species found in mountainous regions across Europe, Asia, Africa, ...
[2]
Caprinae (chamois, goats, serows, sheep) - New Hampshire PBS
Caprinae There are 36 species in this subfamily of medium-sized ungulates (hoofed animals.) They are found in mountainous regions of Asia, Africa, ...Missing: taxonomy | Show results with:taxonomy<|control11|><|separator|>
[3]
Bovids VI: Sheep, Goats, and Relatives (Caprinae) | Encyclopedia.com
The Rupicaprini has four genera and nine species: serows (Capricornis), three species; gorals (Naemorhedus), three species; mountain goat (Oreamnos), one ...
[4]
(PDF) Goats and Sheep: Subfamily Caprinae - ResearchGate
Jun 17, 2024 · The caprinae subfamily contains the sheep, the goats and related species such as the gorals, serows, tahrs and rupicaprids.
[5]
Goat antelope | Habitat, Diet & Adaptations - Britannica
Nov 26, 2008 · Goat antelopes are well adapted to life in precipitous, rocky terrain and can stand the cold well. Males are the same size as or slightly larger than females.
[6]
Wild sheep and goats and their relatives : status survey and ... - IUCN
Wild caprinae, including sheep and goats, are an extremely valuable group of mammals. While most live in mountains, some inhabit desert grasslands, ...Missing: distribution | Show results with:distribution
[7]
Taxonomy browser Taxonomy Browser (Caprinae) - NCBI
Caprinae NCBI BLAST name: even-toed ungulates & whales Rank: subfamily Genetic code: Translation table 1 (Standard) Mitochondrial genetic code:
[8]
Phylogenetic Relationships and Evolutionary History of Goats ...
Aug 13, 2025 · The genus Capra belongs to the subfamily Caprinae within the family Bovidae (Artiodactyla: Mammalia) and it comprises domestic goats as well as ...
[9]
[PDF] New Species of Sinocapra (Bovidae, Caprinae) from the Lower ...
New Species of Sinocapra (Bovidae, Caprinae) from the Lower Pliocene Panaca Formation, ... review in Lindsay et al. 2002). Stock (1921) first recovered and ...
[10]
MAMMALIAN SPECIES 830:1–12 - Capra ibex (Artiodactyla: Bovidae)
May 27, 2009 · Order Artiodactyla, suborder. Ruminantia, infraorder Pecora, family Bovidae, subfamily. Caprinae, tribe Caprini (Simpson 1945). There are ...
[11]
Molecular Systematics of the Subfamily Caprinae (Artiodactyla ...
These differentially weighted parsimony analyses indicate that (1) the subfamily Caprinae is monophyletic, but only with the exclusion of Saiga from the ...Missing: al. | Show results with:al.
[12]
The subfamilies and tribes of the family Bovidae - GENTRY - 1992
Given the primitiveness of Boselaphini and allies, the crucial finding is that Caprinae link with African antelopes and that Antilopini are more remote.
[13]
Biology and Diseases of Ruminants (Sheep, Goats, and Cattle) - PMC
This chapter addresses basic biology, husbandry, and the more common or important diseases of the three ruminant species used most commonly in the laboratory.Missing: Caprinae | Show results with:Caprinae
[14]
https://www.researchgate.net/publication/348917053_Update_on_the_reassessment_of_Caprinae_in_the_IUCN_Red_List
[15]
IUCN SSC Caprinae Specialist Group
Caprinae on the IUCN Red List. read more. CSG assesses species for the IUCN Red List of Threatened Species to inform conservation efforts.
[16]
Update on the reassessment of Caprinae in the IUCN Red List
Jan 31, 2021 · ... genera (goral: six species Naemorhedus spp.; serow: seven species Capricornis spp.) – has been inflated. 2. We have revised the taxonomy and ...
[17]
Molecular evidence for the polyphyly of the genus Hemitragus ...
At least, we propose a new taxonomy, where the three species of tahr are ranged into three monospecific genera: the genus Hemitragus is restricted to the ...
[18]
A new perspective on Ungulate Taxonomy
Aug 19, 2024 · Changes to the standard list of ungulates (covering approximately 250 species) are proposed as new physical and genetic evidence becomes available.
[19]
Bovidae (antelopes, cattle, gazelles, goats, sheep, and relatives)
The subfamily Caprinae consists of goats , sheep , muskox , and relatives. This subfamily of bovids consists of 12 genera, however, the organization of ...Missing: diagnostic | Show results with:diagnostic
[20]
Sexual size dimorphism in domestic goats, sheep, and their wild ...
Nov 23, 2009 · Sexual size dimorphism (SSD) is a widespread phenomenon in different animal taxa, including the subfamily of goats and sheep (Caprinae), ...
[21]
The ruminant digestive system - University of Minnesota Extension
Ruminant stomachs have four compartments: the rumen, the reticulum, the omasum and the abomasum. Rumen microbes ferment feed and produce volatile fatty acids, ...Missing: Caprinae | Show results with:Caprinae
[22]
Microstructure and mechanical properties of different keratinous horns
Jun 6, 2018 · Mammal horns contain a bony core and an exterior keratin sheath. The sheath develops slowly and attains a definitive size and shape by growing ...
[23]
Morphology of the Bones of the Skull in the Sahel Ecotypes of Goats ...
Aug 6, 2025 · Sexual dimorphism is prominent in the frontal bones of goats, since males usually have stronger and larger horns, with a wider horn ...
[24]
Dental Formulas of Various Animal Species - Merck Veterinary Manual
The canine tooth of domestic ruminants has commonly been counted as a fourth incisor. · The maxillary second premolar and maxillary and first molar can be absent ...Missing: Bovidae | Show results with:Bovidae<|separator|>
[25]
Evolution of the sheep coat: the impact of domestication on its ...
Wild sheep and many primitive domesticated breeds have two coats: coarse hairs covering shorter, finer fibres. Both are shed annually.Missing: molting | Show results with:molting
[26]
Mountain goats' winter coats: What can people-powered research ...
Jan 11, 2021 · Mountain goats start shedding coats in May. Males molt earlier than females, and nannies with kids molt later. Molt is also affected by ...Missing: Caprinae woolly coarse seasonal
[27]
The strategies of Alpine ibex to endure harsh over-wintering conditions
Aug 9, 2025 · Decreased endogenous heat production as the major contribution to reduced metabolic rate during winter has been found in many species of ...Missing: underwool | Show results with:underwool
[28]
How Goats Defy Gravity and How it Has Inspired Engineers
Nov 6, 2024 · There are three main features of goats that contribute to their superior balance: optimal centers of mass (COM), highly flexible joints, and specialized hoof ...
[29]
Siberian ibex - Wikipedia
The coat of Siberian ibex is well suited for camouflage in mountains with low or no vegetation. ... Siberian ibexes live mostly above the tree line, in areas of ...
[30]
Too Hot! Goats to the Rescue - by David B. Williams
May 29, 2025 · They accomplish this cold protection by growing guard hairs up to eight inches long and a dense undercoat of twisted, interwoven, one- to two- ...Missing: underwool | Show results with:underwool
[31]
Sheep Eyes vs. Goat Eyes: Is There a Difference? - A-Z Animals
Sep 30, 2025 · Goats have a field of vision between 320° and 340° of their surroundings. Sheep also have good peripheral vision, with a field of view up to 320 ...
[32]
Amazing Goat Eyes and Remarkable Senses!
Dec 21, 2020 · Coupled with their sensitivity to movement, this allows goats to easily spot land predators. ... Goat hearing range and the meaning of bleats.
[33]
Understanding the Ruminant Animal Digestive System
Ruminants are hoofed mammals, including cattle, sheep, and goats, with a unique digestive system that allows them to better use energy from fibrous plant ...Missing: Caprinae key
[34]
The Structural and Functional Capacity of Ruminal and Cecal ...
Similar to ruminal fermentation, in the ruminant's cecum feed digestion is also performed by a specialized consortia of microorganisms. Fermentable ...
[35]
[PDF] Coevolution and Biogeography Among Nematodirinae (Nematoda
The primary physiographic determinant of distribution was the Beringian nexus linking the Palearctic and Nearctic ... In The. Bering land bridge, D. M. Hopkins ( ...<|separator|>
[36]
Goat (feral or wild) | Established pest animal species | Biosecurity
Jan 15, 2025 · Goat (feral or wild) · History of spread. Feral goats have been present in Australia since early European settlement. · Distribution in Victoria.<|separator|>
[37]
Feral goats | Waikato Regional Council
Provide advice and information on feral goat control (Waikato Regional ... Goats (Capra hircus) were introduced to New Zealand in the 1770s. They were ...
[38]
Invasive Animals - Hawaiʻi Volcanoes National Park (U.S. National ...
With the arrival of Captain James Cook also came the arrival of a major invasive species: goats. In 1778, both arrived on the Hawaiian islands and have left a ...
[39]
[PDF] Assessing the levels of intraspecific admixture and interspecific ...
Aug 2, 2021 · Habitat destruction and fragmentation combined with overhunting, climate change, and the introduction of invasive animals and plants have caused ...
[40]
Demography reveals populational expansion of a recently extinct ...
Apr 1, 2021 · Now extinct due to human activities, C. p. pyrenaica was, and still is, one of Europe's most charismatic animals. Recently, interest in this ...
[41]
[PDF] The CIC Caprinae Phenotype System - Regulations.gov
the family Bovidae (cloven hoofed ruminants). Caprinae taxonomy is complicated, often confusing and sometimes controversial. In the past a number or varying ...
[42]
[PDF] Contraints of Winter Habitat Selection by the Mountain Goat in Alaska
During winter in southeast Alaska food availability, predator avoidance and thermoregulation are potentially important constraints on habitat selection by ...
[43]
(PDF) Seasonal Habitat Use and Movements of Mountain Goats ...
Mountain goats in all habitats are strongly associated with escape terrain ... alpine meadows when temperatures increased. Reduced selection for open ...
[44]
Bighorn Sheep and Mountain Goats - SpringerLink
Sep 2, 2023 · Both ungulates inhabit rugged terrain and feed on grasses, forbs, and browse. Bighorn sheep and mountain goats are widely recognized for their ...
[45]
The abundance and persistence of Caprinae populations - PMC
Aug 15, 2022 · Although the 32 extant, autochthonous species of Caprinae have diverse behaviors, body sizes and inhabit different ecosystems, all females birth ...
[46]
Peninsular Desert Bighorn Sheep Behavior
Bighorn sheep are gregarious and their social structure is primarily matrilineal (based on female associations). Females exhibit a high degree of philopatry.
[47]
[PDF] Social Factors Driving Grouping Dynamics in Bighorn Sheep Ewe
Within and amongst groups, bighorn sheep interact with each other, switch groups regularly, and form dominance hierarchies (Festa-Bianchet, 1991; Hass, 1991). ...
[48]
Breeding migrations by bighorn sheep males are driven by mating ...
Mar 6, 2022 · Bighorn sheep Ovis canadensis males establish a linear dominance hierarchy that influences their mating tactics. Some males perform breeding ...
[49]
Correlates and consequences of social status in female bighorn sheep
Social interactions between females were brief in duration and infrequent. Although a dominance hierarchy was evident among the females, it was not linear.Missing: structure | Show results with:structure
[50]
Behavior of Goats - Merck Veterinary Manual
In feral goat populations, group sizes have been observed to range from 1 to 100 individuals. Social dominance is related to horn size in wild and feral goats.Missing: territoriality | Show results with:territoriality
[51]
Behavior of Sheep - Merck Veterinary Manual
Communication between sheep is based on largely visual signals, including stomping, pawing, fleeing, and specific body postures. Individuals in a flock spread ...
[52]
Characteristics of selected behavioural patterns in sheep and goats
Sep 29, 2024 · In goats, social dominance is clearly established and ... Social hierarchy in the domestic goat: Effect on food habits and production.Missing: territoriality | Show results with:territoriality<|separator|>
[53]
Capra nubiana (Nubian ibex) | INFORMATION - Animal Diversity Web
Both sexes exhibit crouching behaviors, tongue-flicking, and scent marking on the tail, beard and chest as part of their reproductive behavior. Reproductive ...
[54]
Social dominance in adult female mountain goats - ScienceDirect.com
Compared with other female ungulates, mountain goat females interacted aggressively much more frequently and their dominance ranks were less stable in time and ...
[55]
The diet of Myotragus balearicus Bate 1909 (Artiodactyla: Caprinae ...
Aug 7, 2025 · The diet of Myotragus must have contained a high percentage of fibers, as evidenced by the contents of petrified droppings (coprolites). These ...
[56]
Desert bighorn sheep in Mexico
We conducted a study of diet composition in Sonora, Mexico from April 1997 to December 1998. We identified 41 plant species consumed by desert bighorn sheep.
[57]
Table: Features of the Reproductive Cycle-Merck Veterinary Manual
Cattle. 10–12 mo, usually first bred at approx. 14–15 mo. Polyestrous all year; 21 d (18–24) ; Sheep. 6–9 mo. Seasonally polyestrous, early autumn to winter; 17 ...Missing: Caprinae parental care
[58]
[PDF] Reproduction Goats - Semantic Scholar
Gestation period. • The medium-wool and meat-type breeds - shorter gestation period than fine-wool breeds. • High temperatures and high nutrition levels may.Missing: Caprinae polyestrous
[59]
Consequences of twinning induction to Noemi ewes by a ...
The pregnancy rate increased in FSH-treated ewes (mean 55%) compared with control (mean 35%). Percent of ewes giving birth to twins was 0, 75, 80, 100, and 50% ...Missing: Caprinae polyestrous
[60]
[PDF] The Behaviour of Sheep and Goats - C. DWYER
As the social group is very important to sheep and goats social isolation is highly stressful, and is more stressful than capture or restraint within the ...
[61]
Weaning lambs - Sheep 201
Early weaning usually means weaning at any time after 21 days of age, but usually before 90 days. Sixty (range 45 to 70) days is a common weaning age in ...Missing: Caprinae | Show results with:Caprinae
[62]
(PDF) Effects of maternal condition, disease status, and behavior on ...
Jan 3, 2025 · In contrast, nutritional condition of females in late winter had a strong, positive effect on summer lamb survival: lambs born to females in ...
[63]
[PDF] Context-Dependent Effects of Nutrition and Dam Behavior on ...
We hypothesized that both spring nutritional condition of ewes and patterns of movement and space use during summer would influence probability of lamb survival ...
[64]
[PDF] The fossil record and evolution of Bovidae: State of the field
The Miocene evolution of Bovidae took place against the backdrop of major climatic and physiographic changes, including the closure of the Tethys Gateway ( ...Missing: divergence | Show results with:divergence
[65]
Molecular Insights into the Evolution of the Family Bovidae
If the molecular association between the Reduncinae and the Caprinae holds, the data at hand would support a Eurasian origin for both the Reduncinae and the ...Missing: debates | Show results with:debates
[66]
(PDF) Molecular phylogeny of caprines (Bovidae, Antilopinae)
Aug 10, 2025 · Caprines include all bovids related to sheep and goat. The composition of the group is controversial and inter-generic relationships have ...
[67]
Phylogenetic Analysis of Snow Sheep (Ovis nivicola) and Closely ...
Wild sheep of the genus Ovis are Holarctic, with a distribution of extant species and subspecies that span two continents. ... Ice Age. Wild Ovis first appeared ...
[68]
New Olonbulukia material and its related assemblage reveal an ...
Dec 26, 2018 · This work reveals an early radiation of stem caprines along the northern side of the rising Tibetan Plateau and indicates a mixed pattern of pan-Tibetan stem ...
[69]
Molecular phylogeny of caprines (Bovidae, Antilopinae): the ...
Dec 21, 2004 · The composition of the group is controversial and inter-generic relationships have been widely debated. Here, we analysed 2469 characters draw ...
[70]
[PDF] Oioceros rothii (Wagner, 1857) from the late Miocene of Pikermi ...
Jun 24, 2002 · Gentry (1970) listed numerous cha- racters on which Oioceros is similar to the. Caprinae. It is not necessary to comment these characters ...
[71]
the case of North American fossil muskoxen Bootherium and Symbos
The musk ox (Ovibos moschatus) is the only surviving member of a group of Pleistocene North American musk ox genera (Praeovibos, Ovibos, Bootherium, ...
[72]
First occurrence of Soergelia (Ovibovini, Bovidae, Mammalia) in the ...
Family Bovidae GRAY, 1821 · Subfamily Caprinae GRAY, 1821 · Tribe Ovibovini GRAY, 1872 · Genus Soergelia SCHAUB, 1951 · Species Soergelia sp. cf. S. minor ...
[73]
Paleoparasitological analysis of the extinct Myotragus balearicus ...
Myotragus balearicus Bate 1909 (Artiodactyla, Caprinae) is an endemic caprine that inhabited the Eastern Balearic or Gymnesic Islands (i.e., Mallorca, Menorca ...
[74]
The late-Quaternary megafauna extinctions: Patterns, causes ...
The extinct species include, for example, a variety of proboscideans and rhinoceroses, giant ground sloths and armadillos, rhinoceros-like toxodonts, and a ...Missing: Caprinae | Show results with:Caprinae
[75]
Caprinae (Bovidae, Artiodactyla, Mammalia) from the Neogene of ...
Bosscha Erdbrink, “Early Samotharium and Early Oioceros from an Uppermost Vindobonian Fossiliferous Pocket at Mardaq near Maraghen in N.W. Iran,” Mitt. Bayer.
[76]
Wild sheep and goats and their relatives : status survey and ...
The main threats to them are over-harvesting, habitat loss and resource competition from livestock. ... This action plan explores the value of caprinae to ...
[77]
Retention of Threatened Status for Argali in Kyrgyzstan, Mongolia ...
May 22, 2002 · Poaching. Local harvest of argali for sport and/or consumption is prohibited. In previous years illegal hunting was acknowledged to be a ...
[78]
(PDF) Recent advances in climate change research related to the ...
Sep 12, 2018 · 2015). As the Caprinae are largely alpine species, it is reasonable to expect that climate change poses a real threat to their conservation, and ...
[79]
Multiple Origins of Foot-and-Mouth Disease Virus Serotype Asia 1 ...
Since 2004, the region affected by outbreaks of this serotype has increased from disease-endemic countries in southern Asia (Afghanistan, India, Iran, Nepal, ...Missing: Caprinae | Show results with:Caprinae