Temporal range: 20–0Ma Early Miocene - Recent
|Images of a few members of the family Bovidae (clockwise from top left) - Sable antelope, sheep, zebu, Chinese goral, nyala and Maxwell's duiker.|
Aepycerotinae (one genus)
The Bovidae are the biological family of cloven-hoofed, ruminant mammals that includes bison, African buffalo, water buffalo, antelopes, gazelles, sheep, goats, muskoxen, and domestic cattle. A member of this family is called a bovid. Consisting of 143 extant species and 300 extinct species, the family Bovidae consists of eight major subfamilies apart from the disputed Peleinae and Pantholopinae. The family evolved some 20 million years ago, in the early Miocene.
All bovids have the similar basic form - a snout with a blunt end, a pair 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.The bovids show great variation in size and pelage colouration. 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 always a pair 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.
- 1 Etymology and taxonomy
- 2 Evolution
- 3 Characteristics
- 4 Ecology and behaviour
- 5 Interaction with humans
- 6 Classification
- 7 References
- 8 External links
Etymology and taxonomy
The name "Bovidae" was given by British zoologist John Edward Gray in 1821. The word "Bovidae" is the combination of the prefix bov- (originating from the New Latin word Bos, meaning an ox) and the suffix -idae. Bovidae is a mammal family placed in the order Artiodactyla (which includes the even-toed ungulates). It includes 143 extant species, accounting for nearly 55% of the ungulates, and 300 extinct species.
Molecular studies have supported monophyly in the family Bovidae (a monophyletic group of organisms comprises an ancestral species and their descendants). The number of subfamilies in the Bovidae is disputed, with suggestions of as many as ten and as less as two subfamilies. However, molecular, morphological and fossil evidence indicates the existence of eight distinct subfamilies : Aepycerotinae (consisting of only the impala), Alcelaphinae (bontebok, hartebeest, wildebeest and relatives), Antilopinae (several antelopes, gazelles and relatives), Bovinae(cattle, buffaloes, bison and other antelopes), Caprinae (goats, sheep, ibex, serows and relatives), Cephalophinae (duikers), Hippotraginae (addax, oryx and relatives) and Reduncinae (reedbuck and kob antelopes). In addition, there are three extinct subfamilies: Hypsodontinae (during the mid-Miocene), Oiocerinae (during the Turolian) and the subfamily of Tethytragus (during mid-Miocene).
In 1992, Alan W. Gentry of the Natural History Museum, London divided the eight major subfamilies of Bovidae into two major clades on the basis of their evolutionary history : the Boodontia, which comprised only the Bovinae; and the Aegodontia, which consisted of the rest of the subfamilies. Boodonts have somewhat primitive teeth, resembling those of oxen; whereas aegodonts have more advanced teeth like those of goats.
There is a controversy about the recognition of Peleinae and Patholopinae, comprising the genera Pelea and Pantholops respectively, as subfamilies. In 2000, American biologist George Schaller and palaeontologist Elisabeth Vrba suggested the inclusion of Pelea in Reduncinae, even though the grey rhebok, the sole species of Pelea, is highly different from the kobs and the reduncines in morphology. Pantholops, earlier classified in Antilopinae, was later placed in its own subfamily Pantholopinae. However, molecular and morphological analysis supports the inclusion of Pantholops in Caprinae.
Early Miocene and before
In the early Miocene, bovids began diverging from the cervids (deer) and giraffids. The earliest bovids, whose presence in Africa and Eurasia in the latter part of early Miocene (20 Mya) has been ascertained, were small animals, somewhat similar to modern gazelles, and probably lived in woodland environments. The Eotragus, the earliest known bovid, weighed 18 kg (40 lb) and was nearly the same in size as the Thompson's gazelle. Early in their evolutionary history, the bovids split into two main clades: Boodontia and Aegodontia. This early split between Boodontia (of Eurasian origin) and Aegodontia (of African origin) has been attributed to the continental divide between these land masses. When these continents were later rejoined, this barrier was removed, and either group expanded into the other's territory. The tribes Bovini and Tragelaphini diverged in the early Miocene. Bovids are known to have reached the Americas in the Pleistocene by crossing the Bering land bridge.
The present genera of Alcelaphinae appeared in the Pliocene. The extinct Alcelaphine genus Paramularius, that was the same in size as the hartebeest, is believed to have come into being in the Pliocene, but went extinct in the middle Pleistocene. Several genera of Hippotraginae are known since the Pliocene and Pleistocene. This subfamily seems to have diverged from Alcelaphinae in the latter part of early Miocene. The Bovinae is believed to have diverged from the rest of the Bovidae in the late Oligocene. The Boselaphini became extinct in Africa in the early Pliocene, and their latest fossils were excavated in Langebaanweg (South Africa) and Lothagam (Kenya).
The middle Miocene marked the spread of the bovids into China and the Indian subcontinent. According to Vrba, the radiation of the subfamily Alcelaphinae began in the latter part of middle Miocene. The Caprinae tribes probably diverged in the early middle Miocene. Caprini emerged in the middle Miocene, and seems to have been replaced by other bovids and cervids in Eurasia. The earliest fossils of the Antilopines are from the middle Miocene, though studies show the existence of the subfamily from the early Miocene. Speciation occurred in the tribe Antilopini during the middle or upper Miocene, mainly in Eurasia. Tribe Neotragini seems to have appeared in Africa by the end of Miocene, and had become widespread by the Pliocene.
By the late Miocene, around 10 Mya, the bovids rapidly diversified, leading to the creation of 70 new genera. This late Miocene radiation was partly because many bovids became adapted to more open, grassland habitat. Aepycerotinae first appeared in the late Miocene, and no significant difference in the sizes of the primitive and modern impala has been noted. Fossils of Obivines, a tribe of Caprinae, in Africa date back to the late Miocene. The earliest Hippotragine fossils date back to the late Miocene, and were excavated from sites like Lothagam and Awash Valley. The first African fossils of the Reduncinae date back to 6-7 Mya. Reduncinae and Peleinae probably diverged in the mid-Miocene.
All bovids have the similar basic form - a snout with a blunt end, a pair 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. Most bovids exhibit sexual dimorphism, with males usually larger as well as heavier than females. All bovids have four toes on each foot – they walk on the central two (the hooves), while the outer two (the dew-claws) are much smaller and rarely touch the ground.
The bovids show great variation in size. On the one hand, the gaur can weigh as heavy as one tonne (1,000 kg (2,200 lb)) and stands 2–3 m (6.6–9.8 ft) high at the shoulder. The water buffalo can be even heavier, and weigh 1,200 kg (2,600 lb); though it is shorter than the gaur, being at most 2 m (6.6 ft). On the other hand, the royal antelope is only 25 cm (9.8 in) tall and weighs at most 3 kg (6.6 lb). The klipspringer is another small antelope, that stands 45–60 cm (18–24 in) at the shoulder and weighs just 10–20 kg (22–44 lb).
There are differences in pelage colouration as well, the colour ranging from a pale white (as in the Arabian oryx) to black (as in the black wildebeest). However, only the intermediate shades, such as brown and reddish brown (as in the reedbuck), are commonly observed. In several species, females and juveniles exhibit a light-coloured coat, while that of males darkens with age. As in the wildebeest, the coat may be marked with prominent or faint stripes. In some species such as the addax, the coat colour can vary with the season. Scent glands and sebaceous glands are often present.
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 always a pair of simple bony protrusions without branches, often having a spiral, twisted or fluted form, each covered in a permanent sheath of keratin. The unique horn structure is the only unambiguous morphological feature of bovids that distinguish them from other pecorans. A study revealed that there was a high correlation between horn morphology and fighting behaviour of the individual. For instance, long horns are intended for wrestling and fencing, whereas curved horns are used in ramming. Another study found that males with horns directed inwards are monogamous and solitary; whereas those with horns directed outwards tend to be polygynous. These results were independent of the body size.
Male horn development has been linked to sexual selection, Horns are small spikes in the monogamous duikers and other small antelopes, whereas in the polygynous they are large and elaborately formed (for example in a spiral structure, as in the giant eland). Thus, to some extent, horns depict the degree of competition among males in a species. On the other hand, the presence of horns in females is likely due to natural selection. The horns of females are usually smaller than those of males, and are sometimes of a different shape. The horns of female bovids are thought to have evolved for defense against predators or to express territoriality, as non-territorial females, which are able to use crypsis for predator defense, often do not have horns.
In bovids, the third and fourth metapodials are combined into the cannon bone. The ulna and fibula are reduced, and fused with the radius and tibia respectively. Long scapulae are present, whereas the clavicles are absent. Being ruminants, the stomach is composed of four chambers :the rumen (80%), the omasum, the reticulum and the abomasum. The ciliates and bacteria of the rumen ferment the complex cellulose into simpler fatty acids, which are then absorbed through the rumen wall. Bovids have a long small intestine; the length of the small intestine in cattle is 29–49 m (95–161 ft). Body temperature keeps fluctuating through the day; for instance, in goats the temperature can change slightly from nearly 37 °C (99 °F; 310 K) in the early morning to 40 °C (104 °F; 313 K) in the afternoon. Temperature is regulated through sweating in cattle, whereas goats use panting for the same. The right lung, consisting of four to five lobes, is around 1.5 times larger than the left, which has three lobes.
Most bovids bear 30 to 32 teeth. While the upper incisors are absent, the upper canines are either reduced or absent. Instead of the upper incisors, bovids have a thick and tough layer of tissue called the dental pad, that provides a surface to grip grasses and foliage. They are hypsodont and selenodont, since the molars and premolars are low-crowned and crescent-shaped cusps. The lower incisors and canines project forward. The incisors are followed by a long toothless gap, known as the diastema. The general dental formula for bovids is 0.0.2-3.3. Most members of the family are herbivorous, however, most duikers are omnivorous. Like other ruminants, bovids have four-chambered stomachs, which allow them to digest plant material, such as grass, that cannot be used by many other animals. Ruminants (and some others like kangaroos, rabbits and termites) are able to use micro-organisms living in their guts to break down cellulose by fermentation.
Ecology and behaviour
The bovids have various methods of social organization and social behaviour, which are classified into solitary and gregarious behaviour. Further, these types may each be divided into territorial and non-territorial behaviour. Small bovids such as the klipspringer, oribi, steenbok are generally solitary and territorial. They hold small territories into which other members of the species, especially are not allowed to enter. These antelopes form monogamous pairs. Many species such as the dik-dik use pheromone secretions from the preorbital glands and sometimes dung as well to mark their territories. The offspring disperse at the time of adolescence, and males need to acquire territories prior to mating. The bushbuck is the only bovid that is both solitary and non-territorial. This antelope hardly displays aggression, and tends to isolate itself or form loose herds, though in a favourable habitat several bushbuck may be found quite close to one another.
Excluding the Cephalophines (duikers), Tragelaphines (spiral-horned antelopes) and the Neotragines, most African bovids are gregarious and territorial. Males are forced to disperse on attaining sexual maturity, and have to form their own territories, while females are not required to do so. Males that do not hold territories form bachelor herds. Competition takes place among males to acquire dominance, and fights tend to be more rigorous in limited rutting seasons. With the exception of migratory males, males generally hold the same territory throughout their lives. In the waterbuck, some male individuals, known as "satellite males", may be allowed into the territories of other males and have to wait till the owner grows old so that they may acquire his territory. Lek mating, where males gather together and competitively display to potential mates, is known to exist among topis, kobs and lechwes.
Interaction with humans
The Bovidae include three of the five domesticated mammals whose use has spread outside their original ranges, namely the cow, sheep and goat; all are from Eurasia. Other large bovids that have been domesticated within the ranges of their wild ancestors are the domestic buffalo (from the Indian water buffalo), yak, zebu (from the gaur or Indian aurochs) and Bali cattle (from the banteng).
Dairy products such as milk, butter 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, while sheep milk is used to make blue Roquefort cheese in France.
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.
- Genus Aepyceros
- Impala, A. melampus
- Genus Aepyceros
- Subtribe Alcelaphina
- Subtribe Damaliscina
- Tribe Antilopini
- Genus Ammodorcas
- Dibatag A. clarkei
- Genus Antidorcas
- Springbok A. marsupialis
- Genus Antilope
- Blackbuck A. cervicapra
- Genus Eudorcas
- Genus Gazella
- G. psolea †
- Arabian gazelle G. arabica †
- Chinkara or Indian gazelle G. benettii
- Queen of Sheba's gazelle G. bilkis †
- Dorcas gazelle G. dorcas
- Mountain gazelle G. gazella
- Saudi gazelle G. saudiya †
- Speke's gazelle G. spekei
- Cuvier's gazelle G. cuvieri
- Rhim gazelle or slender-horned gazelle G. leptoceros
- Goitered gazelle G. subgutturosa
- Genus Litocranius
- Gerenuk L. walleri
- Genus Nanger
- Genus Procapra
- Genus Ammodorcas
- Tribe Saigini
- Genus Saiga
- Saiga S. tatarica
- Genus Saiga
- Tribe Neotragini
- Genus Dorcatragus
- Beira D. megalotis
- Genus Madoqua
- Genus Neotragus
- Genus Oreotragus
- Klipspringer O. oreotragus
- Genus Ourebia
- Oribi O. ourebi
- Genus Raphicerus
- Genus Dorcatragus
- Tribe Boselaphini
- Tribe Bovini
- Genus Bubalus
- Genus Bos
- Genus Pseudoryx
- Saola, P. nghetinhensis
- Genus Syncerus
- African buffalo, S. caffer
- Genus Bison
- Genus Pelorovis †
- Giant buffalo, P. antiquus†
- Tribe Strepsicerotini
- Tribe Ovibovini
- Tribe Caprini
- Genus Ammotragus
- Barbary sheep, A. lervia
- Genus Arabitragus
- Arabian tahr, A. jayakari
- Genus Capra
- Genus Hemitragus
- Himalayan tahr, H. jemlahicus
- Genus Ovis
- Genus Nilgiritragus
- Nilgiri tahr, N. hylocrius
- Genus Pseudois
- Genus Ammotragus
- Tribe Naemorhedini
- Genus Capricornis
- Genus Nemorhaedus
- Genus Oreamnos
- Mountain goat, O. americanus
- Genus Rupicapra
- Genus Cephalophus
- Abbott's duiker, C. spadix
- Ader's duiker, C. adersi
- Bay duiker, C. dorsalis
- Black duiker, C. niger
- Black-fronted duiker, C. nigrifrons
- Brooke's duiker, C. brookei
- Harvey's duiker, C. harveyi
- Jentink's duiker, C. jentinki
- Ogilby's duiker, C. ogilbyi
- Peters's duiker, C. callipygus
- Red-flanked duiker, C. rufilatus
- Red forest duiker, C. natalensis
- Ruwenzori duiker, C. rubidis
- Weyns's duiker, C. weynsi
- White-bellied duiker, C. leucogaster
- White-legged duiker C. crusalbum
- Yellow-backed duiker, C. silvicultor
- Zebra duiker, C. zebra
- Genus Philantomba
- Genus Sylvicapra
- Common duiker, S. grimmia
- Genus Cephalophus
- Genus Pantholops (also classified under Caprinae)
- Tibetan antelope, P. hodgsonii
- Genus Pantholops (also classified under Caprinae)
- Genus Pelea (also classified under Reduncinae)
- Grey rhebok, P. capreolus
- Genus Pelea (also classified under Reduncinae)
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