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Bovids I: Kudus, Buffaloes, and Bison (Bovinae)

Bovids I: Kudus, Buffaloes, and Bison

(Bovinae)

Class Mammalia

Order Artiodactyla

Suborder Ruminantia

Family Bovidae

Subamily Bovinae


Thumbnail description
Small to very large herbivores; males and often females bear unbranched horns on the head; limb structure is typical of the Artiodactyla, with two main toes terminating in hooves; all have a ruminant digestive system

Size
Length 30–170 in (80–435 cm); shoulder height 24–85 in (60–220 cm); weight 36–2,600 lb (17–1,200 kg)

Number of genera, species
9 genera; 24 species

Habitat
Forest, woodland, savanna, scrub, grassland, alpine meadows, prairie, and steppe

Conservation status
Critically Endangered: 2 species; Endangered: 7 species; Vulnerable: 4 species; Lower Risk/Conservation Dependent: 7 species; Lower Risk/Near Threatened: 3 species

Distribution
Africa, Asia, Europe, and North America

Evolution and systematics

The Bovinae is comprised of 24 extant species in three tribes: Boselaphini, Bovini, and Tragelaphini. Genetic evidence supports the idea that the Bovinae is a monophyletic group and a sister group to the subfamily Antilopinae. The tribe Bovini is also monophyletic and a sister group to the Boselaphini. The Boselaphini, today represented by only two living species, the nilgai (Boselaphus tragocamelus) and the chousingha (Tetracerus quadricornis), both of India, are probably the most primitive of the Bovinae and closest to the ancestors of this subfamily. The Bovini tribe includes yak (Bos grunniens), the various species of wild cattle (Bos), the European and American bisons (Bison), Asian (water) buffaloes and anoas (Bubalus), African buffalo (Syncerus caffer), and probably the saola (Pseudoryx nghetinhensis). The yak is sometimes placed into the genus Poephagus, while Bison have been suggested to belong to Bos, as has yak. Yaks appear from behavioral and genetic evidence to be intermediate between cattle and bison. The modern members of the tribe Tragelaphini are all African species, which have probably been separated from other Bovinae for 15 million years. Fossils attributed to nyala (Tragelaphus angasii) have been found in 6.5-million-year-old deposits. The mountain nyala (Tragelaphus buxtoni) is believed to be the precursor of kudu (Tragelaphus strepsiceros and T. imberbis). Other tragelaphines include the bongo (Tragelaphus eurycerus), bushbuck (T. scriptus), sitatunga (T. spekii), and elands (Taurotragus).

Physical characteristics

The smallest species of Bovinae is the chousingha, which weighs 36–45 lb (17–21 kg) and stands around 24 in (60 cm)

at the shoulder. However, most species of Bovinae are medium to large ungulates, ranging up to 2,600 lb (1,200 kg), and several stand over 78 in (2 m) at the shoulder.

Males in all but one species of Bovinae have horns. In most of these species, females have smaller horns. This subfamily includes the only living artiodactyl with more than two horns. Male chousingha, also called the four-horned antelope, possess two pairs of short, sharp horns. Horn shape and relative size vary among species. Tragelaphines have long spiral-shaped horns with smooth surfaces, whereas the wild cattle, African buffalo, and smaller Asian (water) buffalo have shorter, smooth, curved, and often stout horns. Water buffalo have very large curved horns that are often ridged towards the base, while the saola's horns are straight and smooth.

Glands are limited in the Bovinae. The chousingha is the only member of this subfamily with pre-or ant-orbital glands in front of the eyes. Other Bovinae also lack pedal glands.

Most species of tragelaphins have a coat with several thin, white, vertical stripes, and some also have white spots. The contrasting white markings of tragelaphins probably act as disruptive patterns that help camouflage the animal by

breaking up the body outline against the bush and forests they inhabit. In contrast, the eland, although a tragelaphin, is not striped and it inhabits open habitats where disruptive pelage would have little adaptive value. The largest members of the subfamily (e.g., American bison, water buffalo, African buffalo, and members of the genus Bos) have uniformly dark body pelage, although the lower legs of some species are light colored.

Distribution

Bovinae are native to both the New and Old Worlds and range from the north temperate regions south to the tropics. Africa, North America, Eurasia, India, and southern Asia are the main distribution regions. Bovinae are not native to either South America or Australia. The greatest species diversity of Bovinae is found in Africa and southern Asia, with the least in Europe and North America.

Habitat

Depending upon species, Bovinae inhabit a wide variety of habitats, ranging from open grasslands and savannas, thorn and scrubland, to swamps and dense tropical forests. Bovinae can be found at low elevations or above the tree line on the high mountain plateaus of Asia.

Behavior

For most species, males and females generally live apart for most of the year. Adult males live either alone or in all-male groups. Females form groups of varying size, comprised of their young of one to two years age, other females, and sometimes including subadult males. The degree of social grouping varies within Bovinae and is related partly to habitat and to body size. Most tragelaphines, except elands, live solitarily or in small groups. The largest species inhabiting open habitats are highly social, forming large groups, although group size often declines when they occupy more visually dense habitat where group cohesion is more difficult to maintain. When Europeans first traveled across the North

American plains and before their populations were decimated, bison were reported to live in immense herds. These are probably rivaled today only by the migratory herds of wildebeest in the Serengeti of East Africa or of barren-ground caribou in the Arctic tundra of North America. In most species, the adult males form separate all-male groups apart from the females, young, and subadults.

Among males, fighting can occur over attendance at cows in heat and involves charging and ramming their horns together. More often, hierarchical disputes are settled by dominance displays that involve swinging the horns and head actively from the side, presumably to enhance their apparent

size. The defeated subordinate may act like a juvenile by lowering its head and placing its nose beneath the dominant's belly as if it were to suckle, or it may simple run away.

Feeding ecology and diet

Bovinae include grazers, grazer-browsers (mixed feeders), and browsers. Some such as the bushbuck (Tragelaphus scriptus) also eat fruits and seed pods. Tragelaphines are primarily browsers feeding on twigs, leaves, fruits, and new shoots of woody species as well as forbs and sometimes grasses, especially when the latter are newly growing. Larger species such as members of Bos and Bison rely more heavily on grasses and forbs but will browse when fresh young growth of shrubs is available. In the African buffalo, the smaller forest subspecies relies on browse rather than grasses, while the larger Cape buffalo feeds almost exclusively on grasses in the savanna, open bush, and riverine habitats it occupies.

Reproductive biology

All species are polygynous, with males mating with several females, while females tend to mate with only one male per mating season. Single young are most common, although in some species twins are not uncommon. Among the Bovinae, gestation is shortest (7.5–8 months) in chousingha, longest in the water buffalo (10–11 months), and 8–9 months for most other species. Age at first reproduction is generally in the second year (i.e., mate first when 2.5 years old) for the larger species, but occurs during the first year (i.e., mate first when 1.5 years old) for many of the smaller species.

Conservation status

A total of 22 species are listed in the IUCN Red List of Threatened Species. However, listed as Endangered is the linh duong (Pseudonovibos spiralis); this species' existence is questionable as it is suspected of being a hoax. The listings for this subfamily are: Critically Endangered: 2 species; Endangered: 7 species; Vulnerable: 3 species; Lower Risk: 10 species. Excluding the linh duong, 11 of the accepted 21 species, or 52%, are listed in categories of conservation concern.

Most threats to Bovinae come from human activities, including hunting and loss of habitat through the encroachment of agricultural lands and loss of forests for timber harvesting. Increased road access is linked with both the latter activities, which enables hunters to reach previously isolated populations.

Both wisent (Bison bonasus) and American bison (Bison bison) are examples of the extent to which some Bovinae have been driven to the brink of extinction at the hand of humans. Wisent were all but wiped out, leaving just a few in Poland,

while American bison once estimated to number over 60 million at the beginning of the nineteenth century had all but been eradicated through concerted hunting efforts by the end of that century. One of the most important conservation efforts to preserve wisent has been carried out in the Bialowieza Forest in Poland, while for American bison, there have been several centers of bison conservation across the continent.

Significance to humans

All species were hunted, and some such as wisent and bison were especially important for European and North American hunters from Paleolithic to historic times. Today, local peoples still hunt members of the Bovinae for food, and trophy hunters value the species with large horns.

Three species of Bovinae are important domestic species. The most common and widespread is cattle (Bos taurus), first domesticated probably some 6,000 years ago. Domestic cattle are represented by numerous breeds around the world and have been bred for meat, milk, hides, as well as draught animals. They are more suited to temperate than tropical regions, although zebu or humped cattle are well adapted to hot climates. Water buffalo is the next most common domestic Bovinae, and, like its wild form, is an animal best suited to tropical regions with high rainfall. Like domestic cattle, water buffalo are kept for their milk, meat, and hides, and provide power primarily for plowing. The yak is of major importance in the high mountain regions of Tibet, Northern India, Nepal, and Afghanistan, where it is well adapted to elevations above 10,000 ft (2,500 m) above sea level. It is also crossed with domestic cattle to create hybrids that are more suitable to lower mountain elevations. Yak and the hybrids are used as beasts of burden to carry loads and plow fields, but also supply milk and sometimes meat. Their hair is valued and is woven into material used for such purposes as making tents.

Species accounts

List of Species

Yak
African buffalo
Greater kudu
Water buffalo
Aurochs
American bison
Chousingha

Yak

Bos grunniens

taxonomy

Bos grunniens Linnaeus, 1766, boreal Asia.

other common names

French: Yack.

physical characteristics

Body length 94.4–127.9 in (240–325 cm); shoulder height 62.9–80.7 in (160–205 cm); tail length 23.6 in (60 cm); weight males 1,100–2,645 lb (500–1,200 kg), females 660–770 lb (300–350 kg). Moderate sexual dimorphism, with females weighing only about 33% of adult males; stocky, ox-like animals with a broad, low-hung head raising steeply to humped shoulders, which are followed by a lower back and rump. Both sexes have long, simple curved, black horns. In adult males, the horns extend up to 37.4 in (95 cm), whereas those of females normally only attain 19.6 in (50 cm). The pelage of wild yak is black with rusty-brown tints and, sometimes, gray hairs on the muzzle. The domesticated yak varies greatly in color from black to light yellow-brown, with many individuals having mottled white patches over parts of their sides and backs. The guard hair is relatively short on the back; on the sides, it can be up to 27.5 in (70 cm) in length, hanging down to form a fringed cape, which extends far enough to the ground to have the legs appear deceptively short. Their long tail is exceptionally bushy throughout.

distribution

The wild yak occurs on the Tibetan Plateau in northern Xizang Province (Tibet) and western Qinghai Province of China. Its historic range included mountains and plateaus of western China, northern India, Nepal, and parts of Mongolia.

Domesticated yak are distributed more broadly across the highlands of central Asia.

habitat

A species of the high altitudes, it is found on high elevation alpine steppes devoid of trees and bushes, down to elevations of 6,560–16,400 ft (2,000–5,000 m). In late summer, yaks exploit this alpine-tundra biome foraging on the pockets of natural pasture. As snow begins to accumulate during fall in these high elevations, they migrate to windswept areas of shallow snow or to lower elevations where there are greater amounts of accessible vegetation, such as in valleys and on plateaus.

behavior

Yaks form herds, but they are segregated by sex. Female herds comprised of adult females, their calves, and juvenile females and males are typically 6–20 animals, but occasionally more than 100. When males become sexually mature, they leave these female groups and join with older bulls to form all-male herds that are generally 2–5 animals, with some as large as 19. Older bulls are often solitary. When threatened, group members either flee or bunch together and collectively face the predator. In either case, if young are present, they tend to be in the center of the group. During the rut in September, mature bulls join female groups during the four-week breeding season. Males compete for females, and rival males fight by trying to gore each other's flanks.

feeding ecology and diet

During summer, yaks consume a variety of growing grasses and forbs such as wildflowers, and supplement their diet with shrubs and lichen. During winter, they consume the dormant grasses and lichens, including some mosses. Yak make altitudinal migrations to exploit seasonal availability of forage.

reproductive biology

Polygynous. Reproduction is timed to benefit from the relatively short season of plant growth and less inclement weather; they mate in September, and after a nine-month gestation period, the single calf is born in June. Females give birth every second year.

conservation status

Classified as Vulnerable, wild yaks face habitat loss and degradation due to livestock grazing on their natural pastures. These alpine/tundra steppes are low in plant productivity and so competition with livestock is exacerbated. Hunting by local people for meat and hides continue to contribute to extirpation of wild yaks. Besides ecological factors, interbreeding between domestic yak and wild yak may pose additional threats to wild populations.

significance to humans

First domesticated over 4,000 years ago, they have supported human life throughout this time in harsh high elevation environments. They are still important to the society and economies of local peoples in many mountain areas in central Asia. Wild yaks are hunted in some areas for meat, wool, and other products. Domestic herds provide milk, cheese, meat, wool, and hides, as well as draft animals and for transporting goods. Their dung is collected for fuel.


African buffalo

Syncerus caffer

taxonomy

Syncerus caffer (Sparrmann, 1779), Cape of Good Hope, South Africa.

other common names

French: Buffle d'Afrique; German: Kaffernbuffel.

physical characteristics

Body length 82.6–118 in (210–300 cm); shoulder height 53–70 in (135–179 cm); tail length 29.5–43.3 in (75–110 cm); weight 1,100–1,984 lb (500–900 kg). Minor sexual dimorphism in body size, with adult females weighing about 17% less than adult males; the smallest subspecies from dense forests is half the body weight of the plains form. The most notable feature is its large head and broad muzzle. Males have relatively short (up to 59 in [150 cm]) but stout horns that typically extend sideways, first curving down, then up along the distal half of their length. Females also have horns, but these are smaller and narrower in girth than those of males. On older males, the broad bases of the horns abut, forming an almost solid plate across the forehead. The forest-dwelling subspecies have shorter and less curved horns. The pelage is short across the body and varies from black to reddish brown, depending on subspecies, sex, and age class. The forest subspecies is reddish brown. There is a fringe of long hairs on the ears and a short mane. The tail is long, ending in a prominent black tuft of hairs.

distribution

At a broad level, it is distributed in Africa from Guinea to southern Sudan and then south to Angola and eastern South Africa. Within large portions of this range, its populations are confined to nature reserves. Across its geographic range, it inhabits

low to high elevations as long as there are sufficient amounts of suitable habitat. It is particularly abundant in parts of Kenya, Tanzania, Uganda, and Democratic Republic of the Congo.

habitat

Most abundant in savannas and riparian complexes (e.g., swamps and river floodplains), but it also occupies forests, grasslands, and shrublands from plains to mountains. In savannas, it requires large areas of dense grass with thickets or trees for resting cover. Populations that are forest dwelling obtain sufficient cover from the trees and bushes, but must meet their food requirements by frequenting small openings among the forest where ground vegetation such as grasses and forbs are abundant. Visit waterholes and muddy areas where they can drink and also wallow.

behavior

Form large groups comprised of subgroups complexly structured by sex and age. This structure is in part hierarchically maintained by social dominance. Herd size is mediated by broad habitat factors. In forest-dwelling buffalo, groups are generally between 3–12 individuals comprised of females, their calves, and yearlings; in open habitats, herds are larger, are usually 50–500 animals, but occasionally reach up to 3,000. These largest herds lack the cohesion of the smaller typical groups. Adult females, their young, and males up to three years old form relatively stable subgroups within the herd; males older than three years form their own subgroups, while many males older than 10 years are solitary. Together, these subgroups move about within the larger herd, as it moves throughout its home range. During the dry season, some of the all-male subgroups may leave the herd to exploit feeding opportunities in an increasingly nutritionally challenging environment. In most areas, breeding occurs in the rainy season soon after calves are born. Males test the urine of females to determine if they are in estrus; when ready to mate, the cow will stand and allow the bull to mount and copulate.

feeding ecology and diet

Primarily grazers on savannas, consuming vast quantities of grasses. However, the subspecies inhabiting forests include a relatively large amount of shrubs in their diet. Not highly selective feeders, so can acquire the bulk of their forage more easily where there are tall grasses.

reproductive biology

Polygynous. Reproduction is tied closely to the rainy season. The gestation period is approximately 11.5 months. Cows first calve when 4.5–5 years old, producing a single young, although occasionally twins are born. Thereafter, mature cows typically reproduce ever two years. Males do not participate in the rut until they are about seven years old.

conservation status

Classified as Lower Risk/Conservation Dependent. After rinderpest epidemics around the turn of the century, it was greatly reduced across much of its range and extirpated in some regions. Since then, populations have increased and the species has reoccupied much of its former range. However, loss of lower elevation habitat to agriculture has restricted it to nature reserves in many areas.

significance to humans

Hunted by local peoples for meat. As well, it has a reputation for being dangerous and so with its formidable size, this adds to its allure for trophy hunters. Such reputation also makes it undesirable in areas inhabited by humans.


Greater kudu

Tragelaphus strepsiceros

taxonomy

Tragelaphus strepsiceros (Pallas, 1766), Cape of Good Hope, South Africa; or Namibia.

other common names

French: Grand koudou; German: Grosskudu.

physical characteristics

Body length males 76.7–96.4 in (195–245 cm), females 72.8–92.5 in (185–235 cm); shoulder height males 47.2–59 in (120–150 cm), females 47.2–55.1 in (120–140 cm); tail length males 13.7–21.6 in (35–55 cm), females same; weight males 496–694.6 lb (225–315 kg), females 396.8–473.9 lb (180–215 kg). Sexual dimorphism is moderate, with females being are 5% shorter in length than males and weigh 27% less than males. Males have longest horns in all the Bovinae, extending 66 in (168 cm) or longer in a double spiral; females normally lack horns, but occasionally some individuals have very small ones. The general pelage color is brown and there are several thin, widely spaced vertical stripes along the body from shoulders to rump; the number of stripes depends upon the subspecies. Adult males also have a notably grayish neck. There is a pronounced gray mane hanging from the neck and a band of longer, darker hair running along the spine from the neck to the rump, but most prominently over the shoulders. This ridge of hairs can be erected to form a narrow crest outlining the back. The head has a white strip across the rostrum (nose), just below the eyes. The moderately long tail is white beneath with a black tip.

distribution

Found widely distributed in Africa, occurring in southern Chad, northern areas of the Central African Republic, western and eastern Sudan, northeast Uganda, Ethiopia, and Somalia, and then south and southwest to South Africa, Namibia, Angola, and southeastern Democratic Republic of the Congo. The species has been extirpated in many regions of its former range, and the current main stronghold is South Africa with some major areas of representation in East Africa.

habitat

Favors open woodlands with scattered and dense brush, and is found where such vegetation occurs, including plains, rocky hills, and low mountains. It requires brushy thickets for resting cover, and can be found along the wooded banks of dry river courses. Generally, they prefer habitats that provide concealment.

behavior

Females form small herds typically of 6–12 individuals, including young, although some form with up to 20 members, consisting of females, their young, and subadult males. Larger groups up 40 will form temporarily. Mature males join these female groups during the mating season, but otherwise live separately for most of the year, either singly or with other males in rather loose groups consisting of up to 10 males.

feeding ecology and diet

Primarily browsers, consuming leaves and twigs of a great variety of shrubs and trees, including the seed pods of acacias. They will also periodically consume grasses and forbs.

reproductive biology

Polygynous. Reproduction is tied closely to seasonal patterns of rain. Females give birth to a single young after a gestation period that has been estimated to be 7–9 months. Young spend most of the day hidden while their mothers go elsewhere to feed. After about two weeks, young join the herd but continue to hide, mainly at night, for another month.

conservation status

Classified as Lower Risk/Conservation Dependent. Kudu populations were decimated at the beginning of the twentieth century by Rinderpest epidemics. As a result, the species was greatly reduced across much of its range and extirpated in some regions. Since then, it has reoccupied much of its former range in South Africa. However, in East Africa, loss of habitat at lower elevations has restricted the species to certain areas, including many protected parks and reserves.

significance to humans

Hunted by local peoples for meat. Male greater kudu are much sought-after by trophy hunters because of their large impressive horns. It is also a favorite zoo animal because its impressive size, interesting pelage, and unusual horns appeal to visitors.


Water buffalo

Bubalus bubalis

taxonomy

Bubalus bubalis (Linnaeus, 1758), Asia.

other common names

English: Asian buffalo; French: Buffle de l'Inde; German: Wasserbüffel.

physical characteristics

Body length 98.4–118.1 in (250–300 cm); shoulder height 59–74.4 in (150–189 cm); tail length 23.6–39.3 in (60–100 cm); weight 1,543–2,645 lb (700–1,200 kg). Females are slightly smaller and weight about 20% less than males. The largest member of the Bovinae, it is a heavy, bulky animal with disproportionately large feet, whose wide hooves help it avoid sinking too deeply in the mud as it moves about wetlands and swamps. Pelage of wild water buffalo, although somewhat sparse, is dark gray to black; domesticated forms can exhibit a range of coat colors. The relatively long tail ends with a bushy tuft of black hairs. Males have massive crescent-shaped pointed horns about 47.2 in (120 cm) long that are almost triangular in cross-section and with heavy ridges on their surface. With the flattened side of the triangular horn facing upward, the horns extend back, almost parallel with the slope of the face. Females also have horns, which after adjusting for their smaller size, are relatively the same size as in males. Domesticated water buffalo have much smaller bodies, almost one-half the size, and also smaller horns than the wild form.

distribution

In the mid-twentieth century, they occurred in two regions. In one region, they were distributed from central peninsular India, north to Nepal, and east to Bhutan. In the second region, they occurred on the Malay Peninsula and north and east to Vietnam, with some on the north portion of the Island of Borneo. This distribution is now severely contracted and fragmented. Currently, wild populations exist in only a few protected areas in India, Nepal, and Bhutan. Another small population occurs in a wildlife reserve in Thailand.

habitat

Inhabit tropical and subtropical forests and grassland biomes. Within these biomes, they are closely associated with water and surrounding habitats ranging from lowland swamps to forests, woodlands, and grasslands, and swamps along alluvial plains. These riparian habitats are a mix of tall, dense grasslands interspersed with open forests, side streams, and small lakes surrounded by short grasses. This complex of habitat types provides abundant forage, as well as forest or dense thickets for cover, along with water, not only to drink, but also for creating muddy wallows in which they spend long periods of the day partially submerged.

behavior

In early studies made in the Assam region of northwestern India, they formed herds of 10–20 individuals, although up to 100 animals were observed in some groups; these groups were very cohesive. In northern Australia, feral buffalo are abundant and likely exhibit social behavior similar to that expected for wild buffalo in their natural range. Adult females, their young, and sub-adult females form small groups of up to 30 members. Around three years of age, males leave these maternal groups to form all-male groups of up to 10 young bulls. Old males tend to be solitary. Several maternal groups are often loosely organized into a larger herd that together occupy a common home range and may come together nightly. Can frequently be found for long periods during the day immersed in water or lying in muddy wallows.

feeding ecology and diet

Primarily grazers, they consume large quantities of grasses, but also eat herbs, aquatic plants, and other vegetation from among the highly productive grasslands and marshes in their home range.

reproductive biology

Polygynous. After a gestation of 300–340 days, cows give birth to a reddish brown to yellow-brown calf. Normally, only one calf is born, but occasionally females will give birth to twins. They nurse for 6–9 months. The female typically produces one calf every second year. Females can first mate when 1.5 years old. Males usually do not leave maternal herds until three years old. In some areas, mating and birth show little seasonal periodicity. This can result in calves being found at any time of the year within a herd. In other areas where there are seasonal differences to forage supplies, there can be more distinct mating and birthing periods.

conservation status

Classified as Endangered. While domesticated water buffalo are very abundant and distributed well beyond their natural historic range, there are probably fewer than 4,000 wild water buffalo left in the world, most of which are in Assam, India. Within the near future, the species faces a high risk of extinction in the wild. Existing populations are very small and restricted to a few nature reserves, most of which are widely separated from each other. The main threats to the species are from hunting and from encroachment by agriculture and livestock. As well, the wild water buffalo is threatened by hybridization with domesticated forms and by diseases transmitted from domestic livestock.

significance to humans

Offer food, hides, and other products to humans. However, their most significant value to humans began 6,000 to 7,000 years ago when they were first domesticated in China near the mouth of the Yangtze River. One of the most important domesticated species in southern and eastern Asia, they are also important in most other subtropical and tropical parts of the world. Large feral populations are established in places such as Australia. Domestic water buffalo continue to provide, in addition to meat, horn, and hides, milk and butter fat. They also provide low cost, accessible sources of power for plowing fields and transportation of people and their crops.


Aurochs

Bos taurus

taxonomy

Bos taurus Linnaeus, 1758, Poloniaelig (or Uppsala, Sweden, according to Thomas).

other common names

English: Wild cattle, wild ox; French: Aurochs; German: Ur.

physical characteristics

Body length 118 in (300 cm); shoulder height 68.8–72.8 in (175–185 cm); tail length 55 in (140 cm); weight 1,763–2,204 lb (800–1,000 kg). Sexual dimorphism is moderate, with females 20% smaller than males; males had horns, up to 31.5 in (80 cm), that extended sideways and then turned upwards and forwards. Females had notably smaller horns. The legs were somewhat longer than in domestic cattle, and their forequarters were larger than their hindquarters. In northern Europe, the adult males were black-brown with a light streak along the back. This pelage contrasted with a whitish circle around the chin and muzzle. Aurochs were gray-brown in southern Europe and red-brown with a light saddle in Africa.

distribution

The original range of wild aurochs was extensive, stretching from Europe to western Russia, and south to the Middle East and northern Africa. Domesticated breeds are now distributed worldwide, except for Antarctica.

habitat

Primarily a species of open forests and woodlands with grassy openings. In Europe, such habitat provided abundant forage in the form of grasses, forbs, and browse. These natural pastures included wet meadows and, in the Pyrenees, sub-alpine parklands. In North Africa, they occupied more open steppe habitat.

behavior

Groups consisting of adult females with their calves and subadults of both sexes, with adult males living in small all-male groups, except during the mating season.

feeding ecology and diet

Beginning in spring and then throughout the summer, aurochs would probably have fed on grasses and forbs, but also browsed on buds and leaves from shrubs and other low vegetation. In fall, they would likely have consumed acorns where available, but still relied primarily on grasses, forbs, and some browse for most of their energy. In winter, they were reported to live on dry leaves in forests. They probably browsed on shrubs and other plants when grasses were unavailable.

reproductive biology

Polygynous. Historical accounts indicate calves were born in May and June after a gestation of nine months. Females probably

gave birth first as two-year olds, and males would become fully active in mating by about their fourth or fifth year.

conservation status

Aurochs is not listed by the IUCN. The last known representatives of the wild form became extinct in Poland in 1627. However, the species in the form of domestic cattle is currently more abundant and widely distributed than ever before.

significance to humans

Aurochs were probably killed for meat and hides by human hunters. However, after domestication, cattle have provided numerous products as well as sources of draft power and transportation. All of these benefits helped facilitate development of human societies and supported agriculture.


American bison

Bison bison

taxonomy

Bison bison (Linnaeus, 1758), Canadian River valley, New Mexico, United States.

other common names

French: Bison américain; German: Bison.

physical characteristics

Body length males 95.2–125.2 in (242–318 cm), females 68.1–109.4 in (173–278 cm); shoulder height males 65.7–73.2 in (167–186 cm), females 59.8–61.8 in (152–157 cm); tail length males 12.9–35.8 in (33–91 cm), females 11.8–20 in (30–51 cm); weight males 1,199–1,999.5 lb (544–907 kg), females 701–1,201 lb (318–545 kg). High sexual dimorphism, with females 20% shorter in length, but 40% less in weight than males; the largest mammal in North America, it has a broad head that seems to be held low because it is followed by a prominent hump over the shoulders created by elongated spines on the thoracic vertebrae. The hindquarters are much smaller than the forequarters. The legs are short and the tail is medium length, terminating with tuft of black hairs. Pelage is brown to dark brown, with longer hairs on the front and top of the head, along the neck, and onto the shoulders and forequarters, including the forelegs. The hair on the rest of the body is much shorter. The ears are partly hidden among the long fur on the head. Although larger in males, both sexes have a beard of long hairs that extend below the chin. Both sexes also have a fringe of long, dark hair running along the lower margin of the neck as far as the chest to form a noticeable mane. Males have a relatively larger shoulder hump than females and also have longer hair on their heads. Males have short black horns that extend to the side and curve upward, then inward near their sharply pointed tips. Females have horns, but they are more slender, and often shorter and more curved than those of males.

distribution

Historically, they occurred from northwestern Canada to central Canada and then south through much of the United States to northern Mexico. Although there are numerous herds privately managed on ranches and game farms, wild populations are reduced to a few remnant populations and confined to a few parks and reserves in North America.

habitat

Inhabited a variety of ecosystems as long as there was sufficient grassland and meadows on which to graze. In northern areas, they inhabited mixed wood forests and parklands as well as extending into the boreal forests where wet sedge meadows occurred. In more southern regions, they inhabited the long and short grass prairies and parklands across central plains and into the foothills of the Rocky Mountains. Primarily grazers, they relied on grasslands as their primary habitat type, even in northern regions where grasslands and meadows were patchily distributed. Present-day, free-ranging populations occur in areas representative of these primary historic habitats.

behavior

Females, calves, and 1–3-year-old males form mixed groups that may contain one or two older males. Additional adult males may join these groups during the rut. Mixed groups tend to be quite cohesive and with strong hierarchies. The three-year-old and older males form small all-male groups of up to 30 animals, although many adult males also occur alone or in pairs. During the rut, these male groups join with female groups. Herds can grow even larger during seasonal migrations. During these migrations bison may travel over 124 mi (200 km) as the animals move to ranges where there is greater forage and shallower snow. Males are polygynous, but an adult male associates with a single female within the larger group. A typical behavior, both sexes wallow; the animals paw a shallow depression in the ground and roll in it.

feeding ecology and diet

Bulk feeders, they are typically not highly selective in their food habits. They rely on obtaining large amounts of low-quality forage, rather than small amounts of high-quality forage. Feed almost exclusively on grasses and sedges. Occasionally, they consume forbs and browse, but these food types are minor components of their diet. This dependence on grasses tied bison closely to short and long grass prairies characteristic of the central part of the North American continent.

reproductive biology

Mating varies across their geographic range, but mostly takes place during July and August in the southern regions, but extending into September in northern regions. Polygynous. Gestation is on average 285 days, after which they give birth to single calf in spring; twins are very rare. Adult females produce calves usually each year, and generally give birth alone, preferring areas with some taller vegetation for concealment. The calf can run within three hours of birth, and are weaned at 7–12 months. For the first three months of life, they are a reddish brown color that is in marked contrast to the darker pelage of the adults.

conservation status

Classified as Lower Risk/Conservation Dependent. Historically, they ranged across half of the North American and numbered in the millions. Their range and abundance has been as severely diminished. Wild American bison occur only in several national parks and wildlife reserves in Canada and the United States. However, they also occur on numerous ranches and privately held herds; in fact, there are more bison in private ownership than on public protected areas, thus as a species they are not at risk of extinction. Threats to wild bison come mainly from diseases and parasites transmitted from domestic livestock.

significance to humans

At one time, American bison were the most important game animal for indigenous people across the plains of central and western North America. They provided all manner of uses and products from meat, bones for tools, hides for blankets, leather for garments, and sinews for twine. Skins and horns were also used in ceremonies by Aboriginal peoples. Bison raised on ranches provide a select market for wild game meat and in some places they continue to be desired trophies for hunters.


Chousingha

Tetracerus quadricornis

taxonomy

Tetracerus quadricornis (de Blainville, 1816), India.

other common names

English: Four-horned antelope; French: Tetracère; German: Vierhornantiope.

physical characteristics

Body length 31.4–43.3 in (80–110 cm); shoulder height 21.6–25.5 in (55–65 cm); tail length 3.9–5.9 in (10–15 cm); weight 33–55 lb (15–25 kg). Unique among Bovinae because the males have two pairs of horns longer pair is 1.9–4.7 in (5–12 cm), smooth, and black, positioned at the top of the head just anterior to the ears; second pair is much smaller, between 0.7–1.5 in (2–4 cm) long, and located on the forehead well between the orbits. Females are hornless, and they exhibit little sexual dimorphism. Pelage is brown to reddish brown on the back, getting lighter on the sides and changing to white along the abdomen and insides of the legs. The anterior surface of each leg is dark brown. The rostrum and forehead is dark brown to blackish. The outer surfaces of the ears are colored similarly and they have an almost-black rim. The upper lip along the sides is white as is beneath the jaw. This changes to a brown neck that is lighter than the back and sides.

distribution

Occur in thickets and wooded areas across most of India and into Nepal; absent from northeast India and the southern quarter of the Indian Peninsula. Occur in many parks and nature reserves, but are increasingly absent from lands outside these protected areas.

habitat

Occupy a variety of habitats such as dry deciduous, dry deciduous scrub, and southern tropical moist mixed deciduous forest types. Many of these forests occur in hilly terrain, but have some flatter areas providing small grassy openings. They frequent these meadows and other small openings, but never stray far from dense thickets of bushes or bamboo where they quickly retreat when disturbed. These landscapes are generally dissected with streams and small rivers, and they are frequently seen near water.

behavior

Normally found as solitary or in groups of two. During the rut, males are aggressive towards other males. Individuals seem to occupy the same home range year-round.

feeding ecology and diet

Eat a variety of plants and plant parts. As they travel their diverse habitats, they consume leaves of shrubs, shoots, fruit, and grasses.

reproductive biology

Mating occurs over a somewhat protracted breeding season from June through September and coincides with monsoon rainy season. Polygynous. Gestation is 7.5–8 months; most young are born February through March. Adult females give birth to between one to three young.

conservation status

Classified as Vulnerable. Although they are still widely distributed in their historical range, local populations face threats from hunting for meat and loss of habitat to deforestation as well as degradation of habitat due to grazing by livestock. Other conservation concerns are that the areas occupied by chousingha are increasingly becoming isolated as habitat fragmentation proceeds through agricultural practices.

significance to humans

The species has little significance to humans other than its unique curiosity as the only mammal with four horns.

Common name / Scientific name/Other common names Physical characteristics Habitat and behavior Distribution Diet Conservation status
European bison Bison bonasus Spanish: Bisonte europeoShort hair in neck area, pelage is same color as relatives. Horns are well-developed. Average female weight 662–1,190 lb (300–540 kg), male 882–2,028 lb (400–920 kg).Temperate coniferous forest like Bialowieza. For feeding, prefer areas of vegetation at least 20 years old. Remain in large groups during winter and break into male-oriented groups during calving season. Most of life is spent feeding and resting.Extinct except where reintroduced — eastern Poland, western Russia, and Caucasus Mountains.Grasses, mosses, trees, and shrubs.Endangered
Gaur Bos frontalis French: GaurCoat is short, dense, and dark brown. Lower legs white to tan, dewlap under shin extends between front legs. Shoulder hump pronounced in adult males. Horns found on both sexes. Bulging gray-tan ridge connects horns on forehead. Head and body length 98–130 in (250–330 cm), shoulder height 67–87 in (170–220 cm), tail length 27–39 in (70–100 cm), weight 1,543–2,205 lb (700–1,000 kg).Tropical woodlands, but have been largely disturbed. Diurnal, live in groups led by a single male.India; Nepal; Myanmar; Thailand; south Tibet and Yunnan, China; southern Vietnam; Cambodia; and Peninsular Malaysia.Grasses, shoots, and fruit.Vulnerable
Banteng Bos javanicus French: BantengMales are dark chestnut brown, cows and juveniles are reddish brown. Both sexes carry horns. Considered the most beautiful of all wild cattle. Adult male weight 1,400–1,760 lb (635–798 kg), females 1,320–1,500 lb (600–680 kg). Average life in wild is 11 years.Open, dry, deciduous forests. Generally occur in groups of 10–30 individuals.Myanmar, Thailand, and Indochina south to northern Peninsular Malaysia; Java; Borneo; introduced to Australia, Bali Island, Sangihe and Enggano Islands; and domesticated in South-east Asia.Grasses, bamboo, leaves, fruit, and young branches of woody shrubs.Endangered
Kouprey Bos sauveli French: Boeuf gris cambodgien; Spanish: Kouprey, toro cupreyDark brown or black. Body is massive, legs are long, backs are humped.Prefer open deciduous forests, grasslands, wooded grasslands, and patches of closed monsoon forest. Nocturnal.Cambodia, southeast Thailand, southern Laos, and western Vietnam.Primarily grazers, but will also consume fruit.Critically Endangered
Nilgai Boselaphus tragocamelusLarge body with short, smooth horns in males. Gray to brownish gray in males, females and young are brown to orangish. Patches of white on face and below chin. White "beard" or tufts of hair present. Stands 46–60 in (119–150 cm) at the s houlder. Weight 240–275 lb (109–306 kg).Ranges from level ground and thin brush with scattered trees, to cultivated plains. Usually herd in small groups of 10 individuals.Eastern Pakistan and northern India south to Bombay and Mysore; and introduced into Texas, United States.Mixed feeders, preferring browse, short grass, and agricultural crops.Lower Risk/Conservation Dependent
Anoa Bubalus depressicornis French: Anoa des plaines; Spanish: Anoa de ilanuraYoung have thick, yellowish brown, woolly hair. Adults have thick, black skin with white or yellowish white stockings on each foreleg. Sometimes there are blotches of white on throat or nape. Horns are triangular and wrinkled. Weight 198–662 lb (90–300 kg).Lowland forests including secondary formations and swampy areas, along coasts, and also at high elevations in mountainous areas. Aggressive toward humans. Can live up to 30 years.Sulawesi.Grasses, ferns, saplings, palm, ginger, and fallen fruit.Endangered
Bushbuck Tragelaphus scriptusMales have horns that spiral once and are parallel to one another. Both sexes have white spots and stripes, patterns vary geographically. Weight ranges from 88 to 176 lb (40–80 kg), males being larger than females.Forest edges or brushy covers near rivers or streams. Aggressive, nocturnal, good swimmers. Generally solitary, but have been seen in small groups.Southern Mauritania to Ethiopia and southern Somalia, and south to northern Namibia and South Africa.Herbs, leaves, twigs, and flowers.Not threatened
Common name / Scientific name/Other common names Physical characteristics Habitat and behavior Distribution Diet Conservation status
Greater kudu Tragelaphus strepsicerosTallest antelopes, shoulder heights range from 39 to 60 in (100–150 cm). Large horns, body coloration varies from reddish brown to blue-gray, with darkest individuals in southern populations. Six to ten stripes along backs, tails tipped with white undersides. Males have beards. Average weight 265–695 lb (120–315 kg).Areas where adequate cover is provided, including bushes and thickets. Live in temporary groups of one to three individuals.Southern Chad, northern Central African Republic, western and eastern Sudan, north-east Uganda, Ethiopia and Somalia south and southwest to South Africa, Namibia, Angola and southeast Democratic Republic of the Congo (Zaire).Herbs, fruits, vines, flowers, and some new grass.Lower Risk/Conservation Dependent
Mountain nyala Tragelaphus buxtoniCoat is grayish chestnut or sandy gray-brown. Males are larger than females and have spiral-shaped horns. Head and body length 75–102 in (190–260 cm), tail length 7–10 in (20–25cm), adult weight 330–660 lb (150–300 kg).High-altitude woodland, bush, heath, moorland, and valley-bottom grassland. Travels in small groups of two to 13 animals.Ethiopia, east of Rift Valley.Herbs, shrubs, grasses, ferns, and lichens.Threatened

Resources

Books

Estes, R. D. The Behavior Guide to African Mammals. Berkeley, CA: The University of California Press, 1991.

Fries, R., and A. Ruvinsky, eds. The Genetics of Cattle. Oxon, MD: CABI Publishing, 1999.

Grzimek, H. C. B., ed. Grzimek's Animal Life Encyclopedia. New York: Van Nostrand Reinhold Company, 1972.

Haltenorth, T., and H. Diller. A Field Guide to the Mammals of Africa. London: William Collins Sons Co. Ltd., 1980.

2002 IUCN Red List of Threatened Species. Gland, Switzlerand: IUCN, 2002.

Jensen, P., ed. The Ethology of Domestic Animals: An Introductory Text. Oxon, MD: CABI Publishing, 2002.

Kingdon, J. East African Mammals: An Atlas of Evolution in Africa. Vol. III, Part C (Bovids). London and New York: Academic Press, 1982.

Lott, D. F. American Bison: A Natural History. Berkeley: University of California Press, 2002.

Mloszewski, M. J. The Behavior and Ecology of the African Buffalo. Cambridge: Cambridge University Press, 1983.

Nowak, R. M. Walker's Mammals of the World, 5th ed. Baltimore and London: The John Hopkins University Press, 1991.

Schaller, G. B. Wildlife of the Tibetan Steppe. Chicago and London: The University of Chicago Press, 1998.

Sinclair, A. R. E. African Buffalo: A Study of Resource Limitations of Populations. Chicago and London: The University of Chicago Press, 1977.

Vrba, E. S., and G. B. Schaller, eds. Antelopes, Deer and Relatives: Fossil Record, Behavioral Ecology, Systematics, and Conservation. New Haven and London: Yale University Press, 2000

Walther, F. R. Communication and Expression in Hoofed Mammals. Bloomington: Indiana University Press, 1984.

Periodicals

Matthee, C. A., and S. K. Davis. "Molecular Insights into the Evolution of the Family Bovidae: A Nuclear DNA Perspective." Molecular Biology and Evolution 18 (2001): 1220–1230.

Harris, R. B., D. H. Pletscher, C. O. Loggers, and D. J. Miller. "Status and Trends of Tibetan Plateau Mammalian Fauna, Yeniugou, China." Biological Conservation 87 (1999): 13–19.

Meagher, M. "Bison bison." American Society of Mammalogists, Mammalian Species 266 (1986): 1–8.

Schaller, G. B., and W. Liu. "Distribution and Status of Wild Yak, Bos grunniens." Biological Conservation 76 (1996): 1–8.

Tulloch, D. "The Water Buffalo, Bubalus bubalis, in Australia: Grouping and Home Range." Australian Wildlife Research 5 (1978): 327–354.

Other

IUCN. 2002 IUCN Red List of Threatened Species.<http://www.redlist.org>.

Huffman, B. The Ultimate Ungulate Page.<http://www.ultimateungulate.com/>.

David M. Shackleton, PhD

Alton S. Harestad, PhD

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