Bovids VI: Sheep, goats, and relatives

(Caprinae)

Class: Mammalia

Order: Artiodactyla

Suborder: Ruminantia

Family: Bovidae

Subfamily: Caprinae

Thumbnail description
Medium- to large-bodied herbivorous, usually mountain-adapted mammals with either short sharp horns or large and ornate horns

Size
35–94 in (90–240 cm); 44–836 lb (20–380 kg)

Number of genera, species
11 genera; 31 species

Habitat
Mountains, deserts, forest, and tundra

Conservation status
Critically Endangered: 1 species; Endangered: 7 species; Vulnerable: 11 species

Distribution
North America, North Africa, and Eurasia; introduced to New Zealand; domesticated varieties on all continents except Antarctica

Evolution and systematics

The taxonomy of the Caprinae is complex and several arrangements have been proposed. Three tribes are generally recognized: Rupicaprini, Ovibovini, and Caprini. Molecular analysis in 1997 suggested that Tibetan antelope or chiru (Pantholops hodgsonii), previously classified in a tribe of its own or with saiga (Saiga tatarica), was closer to the Caprinae than the Antilopinae and may be a basal member of the Caprinae. As of 2003 its phylogenetic relationship to the rest of the Caprinae had not been definitively assigned. There is more disagreement among taxonomists at species and subspecies levels. IUCN's Caprinae Specialist Group (CSG) adopted a working classification for the Caprinae Action Plan in 1997 that contained 31 species and 81 subspecies. CSG subsequently established a Taxonomy Working Group to examine outstanding problems.

The tribe Ovibovini contains two genera, each with a single species: musk ox (Ovibos moschatus) and takin (Budorcas taxi-color). Musk ox has two listed subspecies and takin has four. The Rupicaprini has four genera and nine species: serows (Capricornis), three species; gorals (Naemorhedus), three species; mountain goat (Oreamnos), one species; and chamois (Rupicapra), two species. Serow (Capricornis sumatraensis) has five subspecies. Some authors include Formosan serow (C. swinhoei) with the Japanese species (C. crispus). Himalayan goral (Naemorhedus goral) has two subspecies; N. baileyi has two, and N. caudatus has four. No subspecies have been identified in Oreamnos. Seven subspecies are listed for northern chamois (R. rupicapra), but by the end of 2002 the validity of all of these had not been confirmed. Southern chamois (R. pyrenaica) has three listed subspecies. The Caprini has five genera and 20 species: Barbary sheep (Ammotragus), one species; tahrs (Hemitragus), three species; blue sheep (Pseudois), two species; true goats (Capra), seven species; and wild sheep (Ovis), five species. Ammotragus has six subspecies, but again the validity of all of these is unconfirmed. No subspecies are recognized in the tahrs. Two subspecies of blue sheep (Pseudois nayaur) are listed. The specific status of dwarf blue sheep (P. schaeferi) has been questioned on the ground that it may be only an ecotype. The taxonomy of Capra is complex. Earlier arrangements placed Alpine ibex, Siberian ibex, Nubian ibex, Walia ibex, and west Caucasian tur (C. caucasica) together in a single species, C. ibex, but they are separated here. Markhor (C. falconeri) has three subspecies and wild goat (C. aegagrus) has four. Spanish ibex (C. pyrenaica) has four known subspecies, two of which are already extinct.

The genus Ovis is even more complex, with a variety of arrangements proposed. All recent authors have recognized the distinctiveness of the larger argalis (O. ammon) from the smaller mouflon and urials. The mouflon and urial group is classified as either one species (O. orientalis) or as two (O. orientalis for the mouflons and western forms, and O. vignei for the eastern urials). Twelve subspecies were recognized in the IUCN's Caprinae Action Plan. It has proved difficult to establish a generally agreed classification of the argalis at the subspecies level, with a large number of forms having been named. Eight are listed in the Caprinae Action Plan. Seven subspecies (one now extinct) are listed for the bighorn sheep (O. canadensis) and four for the snow sheep (Ovis nivicola). Thinhorn sheep (O. dalli) has two subspecies. Techniques of molecular genetic analysis are expected to help validate these forms and further clarify relationships within the subfamily.

The fossil record is incomplete. Musk ox and takin are descendants of a formerly widespread group in Eurasia and both evolved in the late Pliocene or early Pleistocene. Rupicaprini apparently evolved in Asia, with the Pliocene Pachygazella as a possible ancestor, though the earliest fossils of chamois are known only from the late Pleistocene. An ancestral rupicaprine must have crossed the Bering land bridge into North America and given rise to Oreamnos, which first appeared there during the Wisconsin glaciation. There is general agreement that the Caprini evolved from the Rupicaprini.

The genus Tossunnoria from the early Pliocene of China shows characters intermediate between gorals and Caprini and is regarded as the probable ancestor of modern Ovis and Capra. Nadler and colleagues confirmed in 1973 Geist's 1971 hypothesis that the Caprini evolved via two main lineages: one through a Barbary sheep-like form to true sheep and another goat-like line.

Members of the genus Capra first appeared in the mid-Pleistocene, probably from a tahr-like ancestor. From the fossil evidence, its distribution once reached the Atlantic coast of Europe. There are few fossils of Ovis. The earliest ones known were found in China and Europe and date from the Pliocene. The genus evolved in the mountains of Eurasia from where they radiated northwards and northeast, crossing the Bering land bridge into North America. Earliest evidence of Ovis in North America dates from about 100,000 years ago. Most modern Caprini evolved during the Pleistocene, a time when there were 18 major glaciations and Geist suggested in 1989 that Caprini differ from Rupicaprini as a result of adaptation to colder climates and to open rather than closed habitats.

Physical characteristics

The Caprinae are medium- to large-sized herbivores. Sizes range from around 44 lb (20 kg) in goral to 836 lb (380 kg) in adult musk ox. Captive musk ox have attained weights of 1,430 lb (650 kg). In gorals and serows, males and females are around the same size. In other Caprinae species, males average larger than females, this difference being most marked in the Caprini. The general body plan shows a strong, stocky build with powerful limbs that facilitate rapid movement through precipitous terrain. Argali and urial sheep (Ovis ammon and O. orientalis) are exceptions, and have a lighter, more agile build reflecting their reliance on speed to escape from predators. All Caprinae bear horns and in many species, growth rings are present that can be used to age the animal.

Rupicaprini have short, sharp horns that rise straight from the top of the head. These curve slightly backwards in gorals and serows, and are sharply hooked in chamois. Ovibovini have thicker, more massive horns. In the musk ox, they curve downwards and out and in takin curl outwards then back. Caprini have the largest and most developed horn shapes. In Ovis, Pseudois, and Capra, they show a high degree of sexual dimorphism with horns of the females much shorter and slimmer than those of the males. Horns are long and swept back in ibexes, twisted in markhor and curling in Ovis, reaching their most massive in the argalis (O. ammon). The maximum length recorded for the Altai argali is 66 in (169 cm). Hooves are well adapted to grip securely on rock.

Coat color is inconspicuous and many species blend easily into their background. Coloration ranges from white to black through shades of straw-yellow, sandy and brown to deep reddish brown. Male Barbary sheep have long chest ruffs, Himalayan tahrs have a thick ruff on the foreparts; argalis and urial have shorter throat ruffs. Wild goat species have beards of varying length. Tails are relatively short. Light rump patches are well developed in Ovis, but absent in rupicaprids, excepting the highly gregarious chamois (Rupicapra).

Distribution

The subfamily is mainly distributed in the Northern Hemisphere; in North America from the Arctic south to northern Mexico; across the whole of northern Africa and the highlands of Ethiopia; in all the mountains of Europe and Asia, extending to southern India and Sumatra. Musk ox is now restricted to North America and Greenland, but formerly occurred in northern Siberia and may have survived there until early recent times. In historic times, it also occurred much more widely in North America and in Western Europe. The takin is found in the mountains of western China and the eastern Himalaya, west to Bhutan. Rupicaprini have a disjunct distribution, with chamois distributed throughout the mountains of central and southern Europe, Turkey and the Caucasus, and gorals and serows are restricted to Asia. Their combined distribution extends from Pakistan to the Russian Far East and the Korean Peninsula. Japanese serow (Capricornis crispus) and Formosan serow (C. swinhoei) are endemic to Japan and Taiwan, respectively. Oreamnos is found in southeast Alaska and mountains of northwestern Canada and the northwest United States. Serow has an extensive range from the western Himalaya to Gansu and Sichuan provinces in China and through Southeast Asia to Sumatra. Barbary sheep are endemic to northern Africa and once occurred on all ranges and rocky massifs in the Sahara and subdesert zones. Himalayan tahr are distributed in a narrow band along the southern side of the Himalaya from the Indian state of Jammu and Kashmir to central Nepal. Arabian tahr are endemic to the mountains of southeast Arabia and Nilgiri tahr are restricted to a small range in the southern part of the Western Ghats in India. The distribution of Pseudois is centered on the Tibetan Plateau and bordering ranges. Its range extends in the west to the Karakoram Mountains of northern Pakistan and penetrates to the southern side of the Himalaya in a few places. Dwarf blue sheep have a very limited distribution in river gorges of southwest China. The genus Capra is distributed through the mountains of Europe and Asia, from the Alps to Lake Baikal, and in northeast Africa. Wild goat, the progenitor of domestic goats, is found in Crete and a few other Aegean islands, and through the Caucasus, Turkey, and Iran to Afghanistan and Pakistan. Markhor has a relatively limited distribution in Pakistan, Afghanistan, and Tajikistan. Spanish ibex occurs on several ranges in the Iberian Peninsula. West Caucasian tur and east Caucasian tur occupy the west and east of the Caucasus, respectively, with a narrow hybrid zone in the center of the range. Alpine ibex occur throughout the European Alps. Siberian ibex have an extensive distribution through the mountains of Central Asia from the Hindu Kush and Pamirs to the ranges of southern

Siberia. Walia ibex survive in a tiny area of the Simien Mountains of Ethiopia. The genus Ovis occurs from Western Europe across Eurasia to northeastern Siberia and into North America. Mouflons and urial are found on the islands of Corsica and Cyprus and eastwards to the upper Indus Valley in the extreme north of India and in Central Asia. Argalis also have a wide Central Asian distribution. This extends from just south of Lake Baikal in the north southwards to the northern side of the Himalaya, and from Kazakhstan in the west to Inner Mongolia in the east. Snow sheep are restricted to mountain ranges of northeast Siberia. Thin-horn sheep occur in Alaska and northwest Canada, while bighorns have a much more extensive range in western Canada and through the western United States, reaching Baja California in Mexico.

Introductions have extended this original distribution. Himalayan tahr and chamois have been introduced successfully to New Zealand. Barbary sheep are among several species of exotic bovids that have been introduced to the southwest United States for sport hunting. Barbary sheep have also been introduced to Spain, mouflons to several parts of Europe, and muskox to Norway. Feral domestic sheep and goats have established populations in several countries and on many islands.

Habitat

Caprinae are primarily associated with hilly or mountainous terrain, except for the tundra-living muskox. The altitudinal range within which they occur is wide. Nubian ibex have been recorded on cliffs around the Dead Sea down to 1,150 ft (350 m) below sea level. Long-tailed goral in the Russian Far East and snow sheep in Siberia both occur on coastal cliffs at or near sea level. In the Himalaya, Karakoram mountains, and on the Tibetan Plateau, Siberian ibex have been recorded at elevations of 22,000 ft (6,710 m), blue sheep at 21,500 ft (6,500 m), and argali at 19,000 ft (5,800 m).

Caprinae are adapted to a wide range of climatic conditions. Musk ox, thinhorn sheep, and snow sheep inhabit Arctic and subarctic regions year-round. Aoudad, Arabian tahr, Nubian ibex, and desert bighorn have adapted to the extreme desert heat of the Sahara, Arabia, and southwest United States. Urial in the deserts of Central Asia need to cope with high summer temperatures and very cold winters. All these desert-living species can obviously tolerate extreme aridity. Other species such as Nilgiri tahr, takin, and serows in subtropical Asia also live in seasonally wet habitats. Red goral (Naemorhedus baileyi) habitat in southeast Tibet has abundant rainfall, over 78 in (198 cm) annually.

Most species are primarily associated with rugged, precipitous terrain, either open or forested. Mountain goat, Capra species, Barbary sheep, tahrs, blue sheep, snow sheep, and

North American sheep utilize cliffs for escape and occur in steep, rocky, or broken country. Serows, gorals, and takin occupy forest or scrub-covered mountains. Some species seem to revel in the most difficult and inaccessible parts of the mountain ranges. Himalayan tahr, markhor, chamois, and goral move utilize both forests or scrub, and alpine meadows or other open areas. Argali and urial prefer more open and less rocky parts of the ranges but they do not avoid steep terrain.

Vegetation types occupied also show a wide range, from desert and semidesert through all montane zones to the high-cold environments of the Tibetan Plateau. Forested habitats include boreal evergreen, temperate deciduous, bamboo, and rhododendron scrub and subtropical forests of Southeast Asia. Walia ibex occur in giant heath scrub in Ethiopia's Simien Highlands.

Behavior

Caprinae mainly occur singly or in small- to medium-sized groups. Larger groups may form at the rut, at feeding grounds in winter, or occasionally at other times. Herds of up to 500 east Caucasian tur, 400 bharal, 300 takin, and 300 chamois have been reported, but these numbers are not typical. Musk ox, wild goat, and Nilgiri tahr all have a maximum group size of around 100. Serow are mainly solitary. Adult females, young and subadult, or young males often form groups, with adult males remaining separate, either solitary or in small groups, and joining the others for the rut. However, social systems in most species are not clear-cut, and mixed groups, nursery groups, and solitary males may all be seen at the same time of year, as, for example, in the case of the blue sheep.

Senses of sight and hearing are well developed. Alarm calls consist of a variety of sneezes, snorts, whistles, and hisses. Many species also stamp their feet in alarm. Most species are excellent climbers and adept at moving over precipitous terrain, seeking refuge from predators on cliffs. Argali and urial depend on speed for escape. When threatened by predators such as wolves, musk ox form a tight circle with heads lowered and young animals inside the ring.

Home range size varies greatly with species and habitat. Summer and winter ranges are commonly used. Bighorn sheep may use up to five home ranges annually. The maximum distance traversed by bighorns between winter and summer ranges is about 30 mi (48 km). In desert species, the location of water sources is an important influence on use of home ranges. Introduced aoudad in Texas had home ranges of 7.44 mi² (19.25 km²) in summer and 1.02 mi² (2.64 km²) in winter. Serow have been known to make well-marked tracks through their forest ranges. Japanese serow of both sexes mark the boundaries of their range with scent from preorbital glands. Arabian tahr mark their range with scrapes in the ground made by the forefeet and renewed regularly.

A common activity pattern is basically crepuscular, with feeding taking place mainly in the early morning and late afternoon or evening. The day is spent resting in shelter on or near cliffs, in caves, or in dense scrub. Ibex, urial, and blue sheep bed for the night in groups at the top of scree slopes, scraping out a smooth sleeping place and with adult animals facing in both directions to watch for danger.

Most species living in high mountains undertake altitudinal movements to lower elevations in winter to avoid cold temperatures or deep snow and return to higher altitudes in spring. The extent of these movements may be as much as 6,560 ft (2,000 m). Musk ox, in contrast, move to more exposed slopes in winter where winds prevent the buildup of snow.

Feeding ecology and diet

All members of the subfamily are generalist herbivores. A wide range of plant material is eaten: grasses, sedges, herbs, shrubs, buds, shoots, and twigs of trees; fruits, acorns, bark, moss, lichen, and fungi. Long-tailed goral in the Russian Far East eat marine grasses and even seaweed. The Caprinae includes both grazers and browsers and most species are both, to a greater or lesser extent. In a large proportion of the global range of Caprinae, there is a significant difference in forage quality between summer and winter. In winter, animals may be forced to eat dry vegetation, twigs, shrubs, and evergreen shoots with a low nutritional content and that may contain secondary compounds. As a result, they lose condition and may even die of starvation. Many species descend in spring to reach the first flush of green growth. Lush mountain pastures containing grasses and sedges provide rich feeding grounds, and it has been estimated that Siberian ibex in Pakistan could gain up to 44 lb (20 kg) in weight by the end of the summer. Invertebrates, including scorpions and beetles have been found in the stomachs of urial in Turkmenistan.

Blue sheep, Siberian ibex, urial, bighorn sheep, musk ox, and Japanese serow all dig through snow to reach winter forage.

Wild goat species may stand up on their hindlegs to gain access to browse and use a foreleg to pull down the vegetation. Himalayan tahr and markhor have been observed climbing into oak trees to feed on the leaves, with some markhor reaching heights of 19–29 ft (6–9 m) above the ground. Serow and mountain goats are also known to climb into trees growing horizontally out of cliffs to feed. Takin are mainly browsers and they may push over young trees to reach the leaves and shoots.

While water needs may be partially met from vegetation consumed or dew that has condensed on vegetation, most species drink from streams and springs, and those living at high altitudes eat snow in winter. Salt licks are also important to many species: mountain goats, for example, may travel several miles (kilometers) to reach them. Blue sheep have been known to lick urine-soaked areas near human encampments and takin consume soil at certain seasons, possibly for its mineral content.

Reproductive biology

Most Caprinae are polygamous, with dominant males enjoying priority access to females. Dominance is established through displays, threats, and direct combat, prior to or during the rut. Fighting may involve locking horns and twisting, direct head-to-head clashes, or lateral or flank attacks. Head-on fighting may involve two animals running straight at each other before clashing horns, or rising up on their hind legs, then crashing down together. The sound may carry a long way through the mountains and all-out bouts between animals such as argali with their massive horns are an impressive spectacle. Thickened front parts of the skull protect them from damage. Rupicaprini do not use direct head butts but attack the flanks of rivals, attempting to stab with their short horns. Rut-related mortality is reported in musk ox and mountain goat. Male displays to females include tail raising, urine spraying, lip curls, low stretches, and kicks with the foreleg. Chamois males bob their head up and down in front of females and Himalayan tahr also nod and shake their heads in display. Mountain goat males mark vegetation during the rut with glands behind the horns, dig rutting pits, and paw the soil onto their flanks and undersides.

Caprinae living in northern latitudes and at high elevations show a strong seasonality in breeding. The particular period varies with locality and is timed so that births coincide with an abundance of fresh green growth in spring or early summer to meet the nutritional needs of lactation and growth of the young. Cold or wet weather at this time increases juvenile mortality. Walia ibex and Barbary sheep may breed throughout the year, but still show seasonal peaks. Gestation period is around eight months in musk ox and five to seven months in other species. Single young are the norm, but twins are not uncommon in some species.

Pregnant females seek out secluded areas to have their young. Young Caprinae can stand soon after birth but normally hide for two to three days before following the female. Most species are weaned in four to five months.

Conservation status

Nineteen species are considered threatened in the 2002 IUCN Red List. The Walia ibex (Capra walie) is Critically Endangered. Its population is estimated to have decreased from 400 in 1983 to 180 in 1996. Six species are Endangered, including two species of tahr. Nilgiri tahr (Hemitragus hylocrius) and Arabian tahr (H. jayakari) have small populations and limited geographic ranges. West Caucasian tur (Capra caucasica) is restricted to a small area of the western Caucasus, while Nubian ibex (Capra nubiana) and markhor (Capra falconeri) have suffered heavily from indiscriminate hunting. Dwarf blue sheep (Pseudois schaeferi) also has a very restricted distribution in the gorge of the upper Yangtze. Six of the subspecies listed in the 2002 Red List were Critically Endangered, and 15 are Endangered. Serow, Arabian tahr, Walia ibex, and several Caprinae subspecies are listed as Endangered by the U. S. Fish and Wildlife Service.

The greatest recent threat to wild Caprinae has been uncontrolled hunting, a factor that intensified sharply during the twentieth century with the introduction of powerful and accurate modern weapons and improved vehicle transport. Indiscriminate hunting has adversely affected all species, driving several towards extinction, wiping out many small populations, and reducing ranges. Other factors with a negative impact include increasing competition with livestock, loss of habitat, fragmentation of isolated populations, and road building that improves access to remote mountain areas. These must also have caused the loss of genetic diversity in most species, to a greater or lesser extent.

For many species, accurate population estimates and range details have not been established, especially for forest living species, and assessments of conservation status have to be based on partial information. At the end of 2002, Arabian tahr numbered around 2,000 and Nilgiri tahr fewer than 2,500. Strict legal protection and reintroductions in the United States and Canada have halted or reversed declines and increased mountain sheep populations by almost 50% in a quarter of a century. The Japanese government gave the Japanese serow special protected status in 1955, and its numbers increased from 2,000–3,000 at that time to about 100,000 at the end of 2002.

Introductions and reintroductions have also been successful in redressing declining situations, for example, with the muskox. Alpine ibex were hunted out in Europe by 1850, except for one herd in the Gran Paradiso area of Italy. Animals from this source have since been reintroduced to many parts of their former range in Switzerland, France, and Austria, and these are now thriving. Many protected areas contain important populations of Caprinae and some have been established to conserve remnant populations. The surviving population of Walia ibex lives in Simien National Park. Wadi Sareen Tahr Reserve in Oman was established to protect Arabian tahr, and Eravikulam National Park in India contains the largest remaining population of Nilgiri tahr.

Innovative schemes to restore old strip mining sites in Canada have created new habitat for bighorn sheep, leading to dramatic increases in numbers, doubling of body mass in females over a period of 15 years, and big increases in male body sizes, including new record horns. The areas have also been colonized by many other large mammal species, and numbers of nesting birds have risen steadily. Well-managed sport hunting programs have also helped to conserve Caprinae populations.

Significance to humans

All species provided a valuable source of meat, hides, and wool to indigenous peoples. The frequent depiction of ibex and other species in rock drawings attests to their importance to early hunters. In addition to meat, they utilized skins and wool for rugs and clothing, sinew, bones, and horn. Virtually all the body parts of Alpine ibex were greatly valued for medicinal use, and over-exploitation drove the animal to the edge of extinction by the middle of the nineteenth century. Blood and many other body parts of serow and goral are widely used for medicinal purposes in Southeast Asia.

Most species of Caprinae have been hunted for trophies as well as meat, and the large and showy horns of wild Capra and Ovis species being especially prized. Commercial sport hunting continues to be popular and managed sport hunting programs in Mongolia and other Central Asian countries have provided a valuable source of income. The larger horns of wild sheep and goats are used for decoration and a variety of other purposes. In southern Arabia, horns of Nubian ibex are placed on the corners of houses to ward off evil spirits. In the northwest Himalaya, large numbers of horns from both wild and domestic species are placed on village altars and on cairns on top of passes as votive offerings.

Without doubt, the greatest significance of Caprinae for humans lies in their role as the origin of domestic sheep and goats. Wild goats were first domesticated about 2,700 years ago in the Middle East, serving ever since as a valuable source of meat, milk, and leather. Domestic sheep and goats now exist in hundreds of varieties and are found on all continents, except Antarctica. They number in the millions and far exceed populations of all wild Caprinae. They provide a source of meat, milk, wool, and hides for people throughout the world and, especially in the case of sheep, are farmed at commercial levels and their products exported. On the negative side, overgrazing by domestic sheep and goats has been widely blamed for degradation of extensive areas of the Mediterranean region and the Middle East.

Species accounts

Resources

Geist, V. Mountain Sheep: A Study in Behavior and Ecology. Chicago: University of Chicago Press, 1971.

Habibi, K. The Desert Ibex. London: Immel Publishing, 1994.

Heptner, V. G., A. A. Nasimovich, and A. G. Bannikov. Mammals of the Soviet Union. Vol. 1, Ungulates. Moscow: Academy of Sciences, 1961.

Lovari, S., ed. The Biology and Management of Mountain Ungulates. London: Croom Helm, 1985.

Macdonald, D. W., and P. Barrett. Mammals of Britain and Europe. London: Collins, 1993.

Nievergelt, B. Ibexes in an African Environment. Berlin: Springer Verlag, 1981.

Roberts, T. J. The Mammals of Pakistan. London: Ernest Benn, 1977.

Schaller, G. B. Mountain Monarchs. Chicago: Chicago University Press, 1977.

Shackleton, D. M., ed. Wild Sheep and Goats and Their Relatives. Status Survey and Conservation Action Plan for Caprinae. Gland, Switzerland: IUCN, 1997.

Tener, J. S. Muskoxen in Canada: A Biological and Taxonomic Review. Ottawa: Canadian Wildlife Service, 1965.

Fedosenko, A. K., and D. A. Blank. "Capra sibirica." Mammalian Species 675 (2001): 1–13.

Gray, G. G., and C. D. Simpson. "Ammotragus lervia." Mammalian Species 144 (1980): 1–7.

Mallon, D. P. "Status and Conservation of Large Mammals in Ladakh." Biological Conservation 56 (1991): 101–119.

Munton, P. N. "The Ecology of the Arabian Tahr (Hemitragus jayakari Thomas 1894) and a Strategy for the Conservation of the Species." Journal of Oman Studies 8 (1985): 11–48.

Neas, J. F., and R. S. Hoffmann. "Budorcas taxicolor." Mammalian Species 277 (1987): 1–7.

Rideout, C. B., and R. S. Hoffmann. "Oreamnos americanus." Mammalian Species 63 (1975): 1–6.

Schaller, G. B., et al. "Feeding Behavior of Sichuan Takin." Mammalia 50 (1986): 311–322.

Shackleton, D. M. "Ovis canadensis." Mammalian Species 230 (1985): 1–9.

IUCN Species Survival Commission, Caprinae Specialist Group, Marco Festa-Bianchet, Chair. Department of Biology, University of Sherbrooke, Sherbrooke, Quebec J1K 2R1 Canada. Web site:

[Article by: David P. Mallon, PhD]