Bovidae

Bovidae Temporal range: 20–0  Ma PreꞒ Ꞓ O S D C P T J K Pg N Early Miocene present
Example Bovidae (clockwise from top left) – domestic cattle (Bos taurus), mountain gazelle (Gazella gazella), impala (Aepyceros melampus), blue wildebeest (Connochaetes taurinus), and mouflon (Ovis gmelini)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
Infraorder:	Pecora
Superfamily:	Bovoidea
Family:	Bovidae Gray, 1821
Type genus
Bos Linnaeus, 1758
Subfamilies
Aepycerotinae (1 genus) Alcelaphinae (4 genera) Antilopinae (3 tribes and 14 genera) Bovinae (3 tribes and 15 genera) Caprinae (3 tribes and 15 genera) Cephalophinae (3 genera) Hippotraginae (3 genera) Nesotraginae (1 genus) Oreotraginae (1 genus) Reduncinae (3 genera) †Hypsodontinae Alternate taxonomy: Antilopinae (10 tribes and 45 genera) Bovinae (3 tribes and 15 genera) †Hypsodontinae

The Bovidae comprise the

The bovids show great variation in size and pelage colouration. Except some domesticated forms, all male bovids have two or more horns, and in many species, females possess horns, too. The size and shape of the horns vary greatly, but the basic structure is always one or more pairs of simple bony protrusions without branches, often having a spiral, twisted or fluted form, each covered in a permanent sheath of keratin. Most bovids bear 30 to 32 teeth.

Most bovids are

The greatest diversities of bovids occur in

The name "Bovidae" was given by the British zoologist John Edward Gray in 1821.^[1] The word "Bovidae" is the combination of the prefix bov- (originating from Latin bos, "ox", through Late Latin bovinus) and the suffix -idae.^[2]

Taxonomy

The

Ruminantia	Tragulina Tragulidae Pecora Antilocapridae Giraffidae Cervidae Moschidae Bovidae

Molecular studies have supported

Below is a cladogram based on Yang et al., 2013 and Calamari, 2021:[13]^[14][15]

Bovidae

Boodontia (Bovinae) Bovini (bison, buffalo, cattle, etc.) Boselaphini (nilgai and four-horned antelope) Tragelaphini (kudus, nyalas etc.) Aegodontia Aepycerotinae (impala) Nesotraginae (suni and bates's antelope) Antilopinae (gazelles, springbok, dik-dik, royal antelope, saiga, etc.) Cephalophinae (duikers etc.) Oreotraginae (klipspringer) Reduncinae (kobs, reedbucks, waterbucks etc.) Caprinae (chamois, sheep, ibexes, goats, muskox, etc.) Alcelaphinae (hartebeest, topi, wildebeest etc.) Hippotraginae (sable antelopes, oryxes etc.)

Alternatively, all members of the Aegodontia, can be classified within the subfamily Antilopinae, with the individual subfamilies being tribes in this treatment.^[14][15]

Evolution

Early Miocene and before

In the early Miocene, bovids began diverging from the

The present genera of Alcelaphinae appeared in the Pliocene. The extinct Alcelaphine genus Paramularius, which was the same in size as the hartebeest, is believed to have come into being in the Pliocene, but became extinct in the middle Pleistocene.^[6] Several genera of Hippotraginae are known since the Pliocene and Pleistocene. This subfamily appears to have diverged from the Alcelaphinae in the latter part of early Miocene.^[19] The Bovinae are believed to have diverged from the rest of the Bovidae in the early Miocene.^[20] The Boselaphini became extinct in Africa in the early Pliocene; their latest fossils were excavated in Langebaanweg (South Africa) and Lothagam (Kenya).^[21]

Middle Miocene

The middle Miocene marked the spread of the bovids into China and the Indian subcontinent.

All bovids have the similar basic form - a snout with a blunt end, one or more pairs of horns (generally present on males) immediately after the oval or pointed ears, a distinct neck and limbs, and a tail varying in length and bushiness among the species.^[25] Most bovids exhibit sexual dimorphism, with males usually larger as well as heavier than females. Sexual dimorphism is more prominent in medium- to large-sized bovids. All bovids have four toes on each foot – they walk on the central two (the hooves), while the outer two (the dewclaws) are much smaller and rarely touch the ground.^[3]

The bovids show great variation in size: the gaur can weigh more than 1,500 kg (3,300 lb), and stand 2.2 m (87 in) high at the shoulder.^[26] The royal antelope, in sharp contrast, is only 25 cm (9.8 in) tall and weighs at most 3 kg (6.6 lb).^[27] The klipspringer, another small antelope, stands 45–60 cm (18–24 in) at the shoulder and weighs just 10–20 kg (22–44 lb).^[28]

Differences occur in

Some species, such as the gemsbok, sable antelope, and Grant's gazelle, are camouflaged with strongly disruptive facial markings that conceal the highly recognisable eye.^[34] Many species, such as gazelles, may be made to look flat, and hence to blend into the background, by countershading.^[35] The outlines of many bovids are broken up with bold disruptive colouration, the strongly contrasting patterns helping to delay recognition by predators.^[36] However, all the Hippotraginae (including the gemsbok) have pale bodies and faces with conspicuous markings. The zoologist Tim Caro describes this as difficult to explain, but given that the species are diurnal, he suggests that the markings may function in communication. Strongly contrasting leg colouration is common only in the Bovidae, where for example Bos, Ovis, bontebok and gemsbok have white stockings. Again, communication is the likely function.^[33]

Excepting some domesticated forms, all male bovids have horns, and in many species, females, too, possess horns. The size and shape of the horns vary greatly, but the basic structure is a pair of simple bony protrusions without branches, often having a spiral, twisted, or fluted form, each covered in a permanent sheath of keratin. Although horns occur in a single pair on almost all bovid species, there are exceptions such as the

In bovids, the third and fourth

Most bovids bear 30 to 32 teeth.

The bovids have various methods of social organisation and social behaviour, which are classified into solitary and gregarious behaviour. Further, these types may each be divided into territorial and nonterritorial behaviour.

Most bovids are diurnal, although a few such as the buffalo, bushbuck, reedbuck, and grysbok are exceptions. Social activity and feeding usually peak during dawn and dusk. The bovids usually rest before dawn, during midday, and after dark. Grooming is usually by licking with the tongue. Rarely do antelopes roll in mud or dust. Wildebeest and buffalo usually wallow in mud, whereas the hartebeest and topi rub their heads and horns in mud and then smear it over their bodies. Bovids use different forms of vocal, olfactory, and tangible communication. These involve varied postures of neck, head, horns, hair, legs, and ears to convey sexual excitement, emotional state, or alarm. One such expression is the

In the mating season, rutting males bellow to make their presence known to females. Muskoxen roar during male-male fights, and male saigas force air through their noses, producing a roar to deter rival males and attract females. Mothers also use vocal communication to locate their calves if they get separated. During fights over dominance, males tend to display themselves in an erect posture with a level muzzle.^[54][55]

Fighting techniques differ amongst the bovid families and also depend on their build. While the hartebeest fight on knees, others usually fight on all fours. Gazelles of various sizes use different methods of combat. Gazelles usually box, and in serious fights may clash and fence, consisting of hard blows from short range. Ibex, goat and sheep males stand upright and clash into each other downwards. Wildebeest use powerful head butting in aggressive clashes. If horns become entangled, the opponents move in a circular manner to unlock them. Muskoxen will ram into each other at high speeds. As a rule, only two bovids of equal build and level of defence engage in a fight, which is intended to determine the superior of the two. Individuals that are evidently inferior to others would rather flee than fight; for example, immature males do not fight with the mature bulls. Generally, bovids direct their attacks on the opponent's head rather than its body. The S-shaped horns, such as those on the impala, have various sections that help in ramming, holding, and stabbing. Serious fights leading to injury are rare.^[31][54][56]

Diet

Most bovids alternately feed and ruminate throughout the day. While those that feed on concentrate feed and digest in short intervals, the roughage feeders take longer intervals. Only small species such as the duiker browse for a few hours during day or night.^[31] Feeding habits are related to body size; while small bovids forage in dense and closed habitat, larger species feed upon high-fiber vegetation in open grasslands. Subfamilies exhibit different feeding strategies. While Bovinae species graze extensively on fresh grass and diffused forage, Cephalophinae species (with the exception of

Sexuality and reproduction

Most bovids are polygynous. In a few species, individuals are monogamous, resulting in minimal male-male aggression and reduced selection for large body size in males. Thus, sexual dimorphism is almost absent. Females may be slightly larger than males, possibly due to competition among females for the acquisition of territories. This is the case in duikers and other small bovids.^[59][60] The time taken for the attainment of sexual maturity by either sex varies broadly among bovids. Sexual maturity may even precede or follow mating. For instance, the impala males, though sexually mature by a year, can mate only after four years of age.^[61] On the contrary barbary sheep females may give birth to offspring even before they have gained sexual maturity.^[62] The delay in male sexual maturation is more visible in sexually dimorphic species, particularly the reduncines, probably due to competition among males.^[3] For instance, the blue wildebeest females become capable of reproduction within a year or two of birth, while the males become mature only when four years old.^[30]

All bovids mate at least once a year, and smaller species may even mate twice. Mating seasons occur typically during the rainy months for most bovids. As such, breeding might peak twice in the equatorial regions. The sheep and goats exhibit remarkable seasonality of reproduction, in the determination of which the annual cycle of daily

Estrus lasts for at most a day in bovids, with the exception of bovines and tragelaphines. Except for the hartebeest and the topi, all bovids can detect estrus in females by testing the urine using the vomeronasal organ.^[31] Once the male is assured that the female is in estrus, he begins courtship displays; these displays vary greatly from the elaborate marches among gregarious species to the fervent licking of female genitalia among solitary species. Females, initially not receptive, ultimately mates with the male which has achieved dominance over others. Receptiveness is expressed by permission for mounting by the male and setting aside the tail by the female. Copulation generally takes a few seconds.^[31][59]

Gestational period varies among bovids - while duiker gestation ranges from 120 to 150 days, gestation in African buffalo ranges from 300 to 330 days. Usually, a single offspring is born (twins are less frequent), and it is able to stand and run by itself within an hour of birth. In monogamous species, males assist in defending their young, but that is not the case in polygynous species. Most newborn calves remain hidden for a week to two months, regularly nursed by their mothers. In some bovid species, the neonates start following about their mothers immediately or within a few days, as in the impala.^[61] Different bovids have different strategies for the defence of juveniles. For instance, while wildebeest mothers solely defend their young, buffaloes exhibit collective defence. Weaning might occur as early as two months (as in royal antelope) or as late as a year (as in muskox).^[59][60]

Lifespan

Most wild bovids live for 10 to 15 years. Larger species tend to live longer;[3] for instance, American bison can live up to 25 years and gaur up to 30 years. The mean lifespan of domesticated individuals is nearly ten years. For example, domesticated goats have an average lifespan of 12 years. Usually males, mainly in polygynous species, have shorter lifespans than females. This can be attributed to several reasons: early dispersal of young males, aggressive male-male fights, vulnerability to predation (particularly when males are less agile, as in kudu), and malnutrition (being large in size, the male body has high nutritional requirements which may not be satisfied).^[66][67] Richard Despard Estes suggested that females mimic male secondary sexual characteristics like horns to protect their male offspring from dominant males. This feature seems to have been strongly selected to prevent male mortality and imbalanced sex ratios due to attacks by aggressive males and forced dispersal of young males during adolescence.^[68]

Distribution

Most of the diverse bovid species occur in Africa. The maximum concentration is in the

arctic tundra. Several bovid species have been domesticated by human beings. The domestication of goats and sheep began 10 thousand years ago, while cattle were domesticated about 7.5 thousand years ago.^[3][59]

Interaction with humans

Domesticated animals

Further information: Domestication

The

domestic yak (from the wild yak), zebu (from the Indian aurochs), gayal (from the gaur) and Bali cattle (from the banteng).^[59]

Some antelopes have been domesticated including the oryxes, addax, elands and the extinct bubal hartebeest. In Ancient Egypt oryxes, addaxes and bubal hartebeests are depicted in carved walls.

The earliest evidence of cattle domestication is from 8000 BC, suggesting that the process began in Cyprus and the Euphrates basin.^[69]

Animal products

Merino wool

is the most valued, with great fineness and softness.

yoghurt, buttermilk and cheese are manufactured largely from domestic cattle, though the milk of sheep, goat, yak, and buffalo is also used in some parts of the world and for gourmet products. For example, buffalo milk is used to make mozzarella in Italy and gulab jamun dessert in India,^[70] while sheep milk is used to make blue Roquefort cheese in France.^[71]

B vitamins.^[72] Bison meat is lower in fat and cholesterol than beef, but has a higher protein content.^[73]

Bovid

Merino wool is 3–5 in (7.6–12.7 cm) long and very soft. Coarse wools, being durable and resistant to pilling, are used for making tough garments^[75]

and carpets.

drinking vessels since antiquity.^[77]

In human culture

Bovidae have featured in stories since at least the time of

Aesop's fables from Ancient Greece around 600 BC. Fables by Aesop include The Crow and the Sheep, The Frog and the Ox, and The Wolf and the Lamb.^[78] The mythological creature Chimera, depicted as a lion, with the head of a goat arising from its back, and a tail that might end with a snake's head, was one of the offspring of Typhon and Echidna and a sibling of such monsters as Cerberus and the Lernaean Hydra.^[79] The sheep, synonymous with the goat in Chinese mythology, is the eighth animal of the Chinese zodiac, and a symbol of filial piety.^[80]

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External links

• Media related to Bovidae at Wikimedia Commons
• Data related to Bovidae at Wikispecies
• "Bovidae" . Encyclopædia Britannica (11th ed.). 1911.
• "Bovidæ" . Collier's New Encyclopedia. 1921.