Bovids are cloven-hoofed ungulates that constitute the family Bovidae, the largest of all extant families within the order Artiodactyla. They include cattle, buffalos, antelopes, sheep, and goats, among others. Though these mammals are most diverse in sub-Saharan Africa, they are also found in parts of Europe, Asia, and North America. They evolved around 20 million years ago in the Early Miocene Epoch.
Male bovids typically possess two or more unbranched horns, though some females, like klipspringers, bear horns, too. These horns often exist in spiral, twisted, or fluted forms and play a key role in sexual selection during mating.
As ruminants, these mammals have four-chambered stomachs and are primarily obligate herbivores, feeding on leaves and fruits. However, some species, like duikers, also consume insects, lizards, birds, and rodents.
They vary greatly in size. The largest bovid, the gaur (Bos gaurus), weighs over 3,300 lb (1,500 kg) and measures 87 in (2.2 m) in shoulder height. In contrast, the royal antelope, one of the smallest bovids, weighs only 6.6 lb (3 kg) and is 9.8 in (25 cm) in height. Similarly, the klipspringer weighs around 22 to 44 lb (10 to 20 kg) and stands 18 to 24 in (45 to 60 cm) tall.
Most bovids are sexually dimorphic, with the males usually being larger than the females.
All bovids possess a blunt snout, oval or pointed ears, one or multiple pairs of horns (generally on males), distinct neck and limbs, and a tail.
They exhibit a wide variety of pelage coloration, ranging from pale white in the Arabian oryx to black in the black wildebeest. However, most bovids have an intermediate fur color, from reddish-brown to brown. Males of many species darken with age, while the females and juveniles remain pale throughout their lives. In some species, like the addax, coat color varies seasonally.
The coat is often marked by faint or prominent stripes, especially in species like the gemsbok, wildebeest, sable antelope, and Grant’s gazelle, which have disruptive facial markings concealing their eyes. Some, like gazelles, exhibit countershading and easily blend against the background (camouflage), thereby dodging the eyes of their predators. Contrasting coloration is also seen in the legs of some bovids, like cattle, bontebok, and gemsbok, which have white stockings.
These unbranched bony protrusions are generally found in male bovids, though females of some species, like klipspringers, are also horned. They originate from an ossified core called the cornual process (in the front bone of the skull) and are covered by a permanent keratinized sheath. Unlike the antlers of deer, these horns are not shed and mostly grow continuously.
Males with horns directed inwards are generally solitary and monogamous, while those with their horns facing outwards are social and polygynous.
Except in some species, such as Jacob’s sheep and the four-horned antelope, horns occur in a single pair.
Being ruminants, their stomachs are characterized by four chambers: the rumen (which covers about 80% of their stomach), the omasum, the reticulum, and the abomasum. The abomasum represents the true stomach, while the other three chambers are called the false stomachs.
The stomach is followed by an elongated small intestine, which in some bovids, like cattle, is as long as 95 to 161 ft.
They have four toes on each foot, and the line of foot symmetry lies between the third and fourth digits (paraxonic). They usually have reduced lateral toes or dewclaws and walk on the central two, the hooves.
The third and fourth metapodials (metacarpals and metatarsals) are fused to form a long cannon bone. The fibula and ulna are reduced and fused with the tibia and radius bones.
These animals usually have 30 to 32 teeth, lacking upper incisors. The upper canines, too, are either much reduced or absent altogether. Instead, they possess a dental pad, a tough tissue layer that aids in gripping foliage. Both the lower incisors and canines project outward, and a long gap or diastema separates the incisors.
These teeth are both hypsodont (enamel extending past the gum line) and selenodont, in which the molars and premolars are low-crowned and have crescent-shaped cusps.
The dental formula for bovids is 0.0.2-3.3/3.1.3.3.
The family Bovidae derives its name from the combination of the Latin words ‘bos’ (meaning ox) and ‘idae’ (referring to the zoological family). It was coined by the British zoologist John Edward Gray in 1821.
Based on molecular, morphological, and fossil evidence, bovids were initially divided into eight distinct subfamilies: Aepycerotinae, Alcelaphinae, Antilopinae, Bovinae, Caprinae, Cephalophinae, Hippotraginae, and Reduncinae. However, two more subfamilies, Nesotraginae and Oreotraginae, were later added.
Currently, Bovidae comprises 143 extant (accounting for about 55% of all ungulates) and 300 extinct species. A phylogenetic study (2003) by Alexandre Hassanin suggests that the families Bovidae and Moschidae (musk deer) are sister groups to the family Cervidae (true deer).
Although, at present, there are 10 subfamilies, this number remains disputed.
Bovids evolved around 20 million years ago, in the Early Miocene Epoch. During this time, they started to diverge from other ungulate groups, like cervids and giraffids.
The earliest bovids were small, resembling modern gazelles, and lived in woodlands. Members of the genus Eotragus (considered the most primitive) weighed around 40 lb (18 kg), about the same size as Thomson’s gazelle.
During this period, these mammals separated into two main groups geographically: Boodontia in Eurasia and Aegodontia in Africa. This division was caused by the physical separation of the Eurasian and African landmasses. Much later, on rejoining, the two groups spread into each other’s territories.
It was later in the Pleistocene Epoch that these mammals reached the Americas by crossing the Bering land bridge.
During this period, bovids expanded into China and the Indian subcontinent. The subfamily Alcelaphinae began diversifying in the Late Middle Miocene, while the Caprinae tribes had already started to split into distinct groups. Additionally, the tribe Antilopini also began to diversify into various species.
By the Late Miocene, bovids underwent massive diversification, and 70 new genera had emerged.
The subfamily Aepycerotinae first appeared in the Late Miocene. Fossils of the earliest members of the tribe Hippotragine, unearthed from sites like Lothagam and Awash Valley, also date back to this period.
All modern-day members of Alcelaphinae first appeared in the Pliocene. One particular genus, Paramularius, originated in the Pliocene but went extinct in the Middle Pleistocene.
Their highest diversity is found in Africa, with the maximum species richness in the savannas of East Africa. However, they are also found in parts of Europe, Asia, and North America.
Most sheep and goats are restricted to Eurasia, though the Barbary sheep and the ibex are found in Africa. Later, domestic bovids have been introduced globally, including in Australia and South America. The muskox is found only in the Arctic tundra region.
Although these mammals were initially grassland dwellers, they have diversified their ecological niches with time. For instance, springbok and oryx inhabit deserts, while bongo and anoa occupy dense forests. Bohor reedbuck, sitatunga, and lechwe live in riparian and swampy environments, whereas mountain goats and takin survive in very high altitudes. Duikers live in equatorial rainforests. Domesticated bovids, on the other hand, live in farms and other man-made habitats.
Although bovids are obligate herbivores that feed on plant matter, some, like duikers, occasionally supplement their diets with animals, like insects, lizards, birds, and rodents. Large bovids particularly consume fibrous vegetation rich in cellulose and lignin.
While members of the subfamily Bovinae consume fresh grass and other forage, those in the subfamily Cephalophinae are frugivores, primarily feeding on fruits. Bovids of the tribes Alcelaphini, Hippotragini, and Reduncini include high proportions of monocots, while those in Tragelaphini and Neotragini (except those in the genus Ourebia) feed on dicots.
Most bovids, except the buffalo, bushbuck, reedbuck, and grysbok, are diurnal. All social activities and foraging peak during dawn and dusk, while they generally rest during mid-day and at night.
Bovids are either solitary or gregarious and engage in a range of social behaviors to communicate with their conspecifics. Many small bovids, like the klipspringer, oribi, and steenbok, are usually solitary, monogamous, and territorial, maintaining small territories of their own. Some species, like the dik-dik, release pheromones from their preorbital glands to mark their territories. The bushbuck, on the other hand, is the only bovid that is solitary but not territorial. It typically isolates itself from the crowd or forms loose herds, though many of these antelopes are often found close to each other.
All African bovids, except duikers, spiral-horned antelopes, and neotragines, are gregarious and territorial. The males disperse after attaining sexual maturity and build their territories. Those that do not form territories form bachelor herds.
Though gregarious, cattle, sheep, and goats are not territorial, and the males, instead of having to maintain territories, invest years in body growth to establish their physical dominance over others.
Bovids typically communicate through various sensory forms, like vocal, olfactory, and tactile cues. Different postures of their body parts, such as neck, head, horns, hair, legs, and ears, convey different states, like alarm or sexual excitement. One such response is the flehmen response, where the animal curls its upper lip, exposing its front teeth and gum, followed by rapid inhalation with its nostrils closed. Through this response, the bovid senses odor, taste, and smell.
They mostly alternate between feeding and ruminating all day. Small bovids, like duikers, forage for a few hours either in the day or at night and prefer browsing dense and closed habitats. Large bovids, on the other hand, browse in open grasslands.
Members of different subfamilies have unique feeding strategies. For instance, those in Antilopinae browse and forage for scattered food resources. In contrast, members of Bovinae both graze on grass and also supplement their diet with scattered food matter.
In addition, those in the family Caprinae are generalists and are flexible feeders foraging in low-productivity habitats, whereas Hippotraginae species graze in arid areas where food supplies are often scarce and unpredictable.
When a predator is nearby, bovids may roar or grunt to warn it and alert other individuals of their group. They also stand motionless, with their head held high with an intent stare.
Some, like gazelles, exhibit stotting (also called pronking) by stiffening their legs and leaping high on them. This response indicates that the gazelle has spotted the predator and is capable enough to put up a fight.
If the predator is too close, they directly attack them by using their S-shaped horns for stabbing or ramming.
Most bovids live for 10 to 15 years, with larger species usually living longer. For example, the bison survives up to 25 years, while the gaur lives for as many as 30 years. In contrast, domestic bovids have an average lifespan of around 10 years.
In polygynous bovids, the females usually live longer than their male counterparts.
Most bovids are polygynous, with a single male mating with multiple females in a single breeding season. However, some, like duikers and other small bovids, are monogamous, and the females are larger than the males, possibly to equip them for female-female combats.
All bovids mate at least once a year, usually in the rainy season or in fall, though smaller species may also mate twice. Bovids, like sheep and goats, are short-day breeders, and their breeding is seasonal and largely influenced by photoperiod.
In most female bovids, the estrus phase (reproductively active phase) lasts for at most a day. All males (except the hartebeest and the topi) detect this phase by smelling the urine of the females using their vomeronasal organs. Once the male is sure the female is in estrus, he begins courtship displays.
During the rut (the mating season of these mammals), the males vocalize to attract potential mates. For example, muskox males bellow and roar to assert their presence to the females. Lekking, where the males gather and engage in competitive displays for females, is observed in topis, kobs, and lechwes.
They exhibit a variety of displays to grab their attention, ranging from elaborate marches (in gregarious species) to licking of female genitalia (in solitary species)
These males compete with each other (male-male competition) for access to females and usually display themselves in an erect posture to establish their dominance. Typically, two individuals of similar physiques contest each other to determine their superiority. Immature and weak individuals usually avoid battling and flee from the ground.
Different bovid families undertake different fighting techniques. For instance, muskoxen and saiga males roar ferociously at other males, while the hartebeest usually fights on all four knees. Gazelles typically box, clash, and fence, landing hard blows on their competitors from a short range. Wildebeests, on the other hand, aggressively butt their heads with their rivals and make circular moves to free their entangled horns. Goats, ibex, and sheep stand upright and hurl onto each other, trying to bring the other down.
Once a male has established its dominance, the females usually mate with the male, though initially, she is reluctant to mate.
Once the female selects the male, she moves her tail aside and signals her receptiveness to him. The pair then copulates, usually taking a few seconds in the process.
The gestation period, ranging from 120 to 150 days in duikers to around 300 days in African buffaloes, is followed by the birth of a single offspring, though twins are born rarely. The offspring is usually capable of standing and running within an hour of birth. They mostly remain hidden for about a week to two months, religiously nursed by their mothers. In monogamous species, the fathers assist in nursing, as well. However, in some bovids, like the impala, the newborn calves follow their mothers immediately or within a few days after birth.
The young are weaned as early as two months in some species, like the royal antelope, while others, such as the muskox, are weaned after a year.
Different bovids attain sexual maturity at different stages of their lives, with some becoming mature immediately after mating. For example, impala males, though mature by a year, mate only after they are four, whereas barbary sheep females give birth even before they have gained maturity.
In Eastern and Southern Africa, large bovids are preyed upon by lions, leopards, cheetahs, and African wild dogs, whereas spotted hyenas, Nile crocodiles, and side-striped jackals attack the smaller species. Similarly, in North America, gray wolves, brown bears, and cougars are their major predators.
In Asia, gray wolves, tigers, dholes, and mugger crocodiles kill these mammals. Moreover, instances of Komodo dragons consuming goats and water buffaloes are also common. Sometimes, even wild dogs and cats consume domestic goats, sheep, and cattle.
Owing to the proper protection of their natural habitats, 67 of the 143 species of bovids currently belong to the Least Concern (LC) category of the IUCN Red List. However, in some regions, they are still hunted for their meat, and overgrazing by domestic animals causes fragmentation of their habitats. As of 2009, aurochs, Queen of Sheebas gazelle, Saudi gazelle, and bluebuck, are extinct in the wild. The Scimitar-horned oryx is also extinct in the wild, being found only in restricted areas like zoos.
The Saola antelope and bighorn sheep are classified as Endangered (EN), while eight other species are listed as Critically Endangered (CR). Additionally, 21 species are considered Vulnerable (VU), and 16 species are Near Threatened (NT). Furthermore, CITES includes 71 species in Appendix I and only one species in Appendix II.
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