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Reproduction

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Most bovids are polygynous, and in some of these species males exhibit delayed maturation. For example, male blue gnus do not reach sexual maturity until 4 years of age, while females become reproductively active between 1.5 to 2.5 years of age. Sexual dimorphism is more prevalent in medium to large bovid species, particularly in members of the subfamily Reduncinae. In general, males of sexually dimorphic artiodactyls become sexually active later in life than females, which is probably due to male-male competition for mates. In some species, males may fight for and defend territory, which gives them breeding rights to females residing within each territory. It is not uncommon for territorial males to try and prevent resident females from leaving (e.g., impalas). Alternatively, males of other species fight for and defend small groups of females known as harems. Adult males that successfully defend their harem often breed with each member of the group, therefore increasing there reproductive fitness. Some bovid species also form leks, a small collection of males that compete for territory or mating rights. Successful males win occupation rights to high quality habitats and thus are able to mate with a greater number of high quality females. Once an individual gains territorial rights, individuals guard their territory and the females within. For example, waterbuck males defend areas of less than 0.5 km2, puka maintain areas of less than 0.1 km2, and lechwe and Uganda kob guard areas of about 15 to 30 m^2. Some species live in large groups consisting of both males and females in which males compete for mating opportunities (e.g., water buffalo). This behavior is somewhat common among members of the subfamily Hippotraginae.

In addition to polygynous mating systems, some species of bovid are monogamous, and male-male competition for mates is less common in these species. As a result, there is decreased selection for large males leading to little or no sexual dimorphism in monogamous bovids. For example, female dik-diks, are solitary and maintain large territories. Thus, male dik-diks are physically unable to defend more than one mate at a time resulting in monogamy. Unless there is a surplus of unmated males, male-male competition is unlikely leading to monomorphism between genders. In fact, females are slightly larger in some monogamous bovids (e.g., duikers and dwarf antelopes), which is probably the result of competition for high quality territories in which to raise their young.

With the exception of hartebeests and topi, all bovids can detect estrus in females. Males sample the urine of potential mates, and high levels of sex hormones in the urine signal that a female is approaching estrus. Males then proceed with courtship behavior in an attempt to secure a mate. Typically, courtship begins with foreleg kicking, chest pressing and finally mounting. Females usually stand to be mounted only at peak estrus.

Mating System: monogamous ; polygynous ; polygynandrous (promiscuous) ; cooperative breeder

Bovids generally breed during fall or the rainy season. Estrus is generally short, usually lasting for less than a couple of days but is longer in non-territorial species. Bovids give birth to a single calf after a relatively long gestation compared to other mammalian families. For example, duiker gestation ranges from 120 to 150 days, while gestation in African buffalo ranges from 300 to 330 days. Calves are usually born synchronously each year during spring, when forage resources are abundant. Adult females reenter estrus within one to two months of parturition. Known as a tending bond, males of non-territorial species often form temporary, exclusive bonds with individual females. Gestation in bovids ranges from 6 months in smaller species to 8 or 9 months in larger species, and some smaller bovids can reproduce biannually. Usually a singe well-developed, precocial calf is born, but twins are not uncommon. Average birth weights vary depending on species. For example, dik-dik calves weigh between 0.5 and 0.8 kg with the males occupying the higher end of the spectrum. New-born eland antelope weigh between 23 and 31 kg. In many gregarious species, young are able to stand and run within one hour of birth.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Like all eutherian mammals, bovids are placental mammals and feed their young with milk. As a result, females are obligated to provide parental care. In polygynous bovids, females provide all parental care without aid from males. In monogamous bovids such as dwarf antelopes, males often defend their young. Weaning may occur as early as 2 months after birth (royal antelope) or as late as one year old as in musk ox.

As calves, bovids can be classified either as hiders or followers. In hider species, mothers hide their young, during which time the mother is typically foraging nearby and on guard for potential predators. Hider mothers return to their calf several times a day for nursing. After nursing, the calf finds a new hiding place nearby. If the species is also gregarious, calves run ahead of their mother during herd movements and hide until their mother has passed. Calves then run ahead and hide again. Mothers with calves of similar age may form mother herds of 2-10 females which continues until the calf is one week to two months old, depending on the species. In follower species young join the herd either immediately or within two days of birth. Newborn wildebeest calves cling to their mother's side and the pair joins a nursery group within the larger herd. Female impalas leave the herd to give birth and rejoin in 1 to 2 days with their young. Upon returning, calves form small nursery groups, which are then guarded by herd females. Some species exhibit group or herd defense of young calves. Males and females alike encircle herd calves, thus protecting them from approaching predators. In many gregarious species, females remain in the herd while males often disperse after independence.

Parental Investment: precocial ; male parental care ; female parental care ; pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Behavior

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Members of the family Bovidae communicate in a number of different ways. Some species are vocal, while others communicate via different body postures and displays. Although vocal communication is limited, during mating season mature males may bellow or roar to intimidate each other and to make their presence known to females. Muskox frequently roar during male-male contests and hold a unique posture that maximizes the intensity of their roar. The ventrorostral ventricle, a vocal ligament that transforms into a large fat pad during maturation, increases the amplitude of the bellow by adding additional resonance space and by directing the sound through a unique pulsing structure. The posture of the male effects how his roar is delivered. Other bovids utilize their nasal passages to roar. Male saiga contract and extend their peculiar noses while forcing air through their nostrils to produce a roaring sound, which is used to deter rival males and attract females. Vocal communication between calves and their mothers help them recognize and locate each other when separated.

In addition to communication that is used to increase reproductive success and offspring survival, bovids also vocalize in an attempt to ward of potential predators. Grunting and roaring, much like those used by competing males, are used to drive off predators and warn herd members. Domesticated bovids are known to vocalize in anticipation of food and native Korean cows vocalize before being fed.

Unlike primates and many carnivorous mammals, bovids are fairly limited in their ability to convey information via facial expressions, thus they rely heavily on postural displays to communicate their intentions. When attempting to communicate dominance or aggression towards competitors or lower ranking individuals, most bovids make themselves look as large as possible. Slow rigid movement and occasionally posing in an erect posture with a level muzzle, is used to exhibit dominance over others. Common aggressive displays include mimic fighting, staring, or shaking their heads wildly to communicate they feel threatened and are ready to fight. Submissive communication includes a lowering of the head or raising the chin so horns rest along the top of the neck. When threatened, bovids often remain still. In some antelope, like impala, lesser kudu, and common eland, individuals may jump in place to signal a potential threat to conspecifics.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Conservation Status

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Currently, many bovid species enjoy sufficient numbers to ensure their survival for years to come. The ICUN red list of threatened species considers 67 of the 143 species listed as “least concern.” This is in part due to the protection of large tracts of land that help offset the detrimental effects of habitat loss. For instance, wildebeest and gazelles in the African Serengeti were fewer than 500,000 during the 1960’s, but had grown to more than a million by the 1990’s. Parks like Serengeti National Park provide ecotourism opportunities and serve as a significant source of income to local economies. As a result, ecotourism enhances the monetary value of wildlife in these countries. In some areas, however, bovids continue to be over exploited for meat and habitat loss due to overgrazing by domestic species, farming, and logging is a significant threat to the persistence of many species. Bovids with limited range and unique habitat requirements are even more at risk. As of 2009, four species of bovid have gone extinct in the wild: aurochs, Queen of Sheebas gazelle, Saudi gazelle, and bluebuck. Scimitar-horned oryx is extinct in the wild and now lives only in zoos. Eight others species are "critically endangered". Saola antelope and bighorn sheep are listed as "endangered". Another 21 species are listed as vulnerable and 16 species are considered "near threatened". CITES, the Convention on International Trade in Endangered Species of Wild Fauna and Flora, lists 71 species under appendix 1 and 1 species under appendix 2.

Bovids are an important food sources for a number of different carnivores. As bovid populations decline, so too will those animals that depend on them. For example, the decline of cheetahs is often attributed habitat loss. However, cheetahs primarily prey upon small to medium sized bovids, specifically gazelle. According to the IUCN Red List of Threatened Species, 2 species of gazelle are extinct, while 10 more are listed as vulnerable, endangered or critically endangered. In north Africa, as preferred prey species have declined, more and more cheetahs are turning to livestock for prey. Consequently, these cheetahs are then killed as pests. As a result, one of the major directives for cheetah conservation is restoration of wild prey species, most of which are small to medium-sized bovids.

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Comprehensive Description

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Bovids are the largest of 10 extant families within Artiodactyla, consisting of more than 140 extant and 300 extinct species. Designation of subfamilies within Bovidae has been controversial and many experts disagree about whether Bovidae is monophyletic or not. While as many as 10 and as few as 5 subfamilies have been suggested, the intersection of molecular, morphological, and fossil evidence suggests 8 distinct subfamilies: Aepycerotinae (impalas), Alcelaphinae (bonteboks, hartebeest, wildebeest, and relatives), Antilopinae (antelopes, dik-diks, gazelles, and relatives), Bovinae (bison, buffalos, cattle, and relatives), Caprinae (chamois, goats, serows, sheep, and relatives), Cephalophinae (duikers), Hippotraginae (addax, oryxes, roan antelopes, sable antelopes, and relatives), and Reduncinae (reedbucks, waterbucks, and relatives). Wild bovids can be found throughout Africa, much of Europe, Asia, and North America and characteristically inhabit grasslands. Their dentition, unguligrade limb morphology, and gastrointestinal specialization likely evolved as a result of their grazing lifestyle. All bovids have four-chambered, ruminating stomachs and at least one pair of horns, which are generally present on both sexes.

Species in the subfamily Bovinae are native to Africa, North America, Eurasia, India, and southern Asia. Bovinae is generally considered to include 24 species from 8 different genera, including nilgai, four-horned antelope, wild cattle, bison, Asian buffalo, African buffalo, and kudu. Sexual dimorphism is highly prevalent in this subfamily, with the males of some species weighing nearly twice as much as their female counterparts. Bovines have played an important role in the cultural evolution of humans, as numerous species within this subfamily have been domesticated for subsistence purposes.

The subfamily Antilopinae includes antelopes, dik-diks, gazelles, and relatives. Small to medium-sized, cover-dependent antelope are found throughout a majority of Africa but occur in particularly high densities in east Africa. Dwarf antelope, steenboks, and dik-diks occur in a variety of different habitats but are also restricted to the continent of Africa. Finally, true gazelles include the genera Eudorcas, Gazella, Nanger, and Procapra, among others. In general, bovids within the subfamily Antilocapinae occur throughout much of Asia and Africa.

Bovids within the subfamily Reduncinae are primarily distributed throughout parts of Eurasia and Africa. Reduncinae is comprised of only three genera, including Redunca (reedbucks), Pelea (rhebok), and Kobus (waterbucks). Species in Reduncinae are medium to large-sized grazers that often have strong ties to water. They also have long hair, and all species exhibit sexual dimorphism, as horns are only present in males.

Bovids in the subfamily Hippotraginae consist primarily of large grazing antelopes with large horns. Hippotraginae species are restricted to Africa and middle-east Asia and are primarily grazers. Most species in this subfamily live in arid habitats and have an erect mane along the nape of the neck. Recent accounts include 8 species from 3 different genera.

Ancelaphinae, consisting of 10 species from 4 genera, includes bonteboks, hartebeest, wildebeest, and relatives. All of the species in this subfamily are nomadic grazers that are native to Africa. Most species are size-dimorphic, with males being 10 to 20% larger than females, and both males and females possess double-curved horns, also known as lyrate.

The subfamily Caprinae consists of goats, sheep, muskox, and relatives. This subfamily of bovids consists of 12 genera, however, the organization of Caprinae is complex and several classifications have been suggested. The International Union for Conservation of Nature (IUCN) currently has a Taxonomy Working Group within their Caprinae Specialist Group to help alleviate some of the outstanding issues within Caprina taxonomy. Caprinids are especially adapted to montane and alpine environments, which explains why this is the only subfamily that is more diverse in Eurasia than Africa. In general, both genders have horns, however, horn morphology in many species is sexually dimorphic.

The subfamily Aepycerotinae consists a single species, the imapala. Aepycerotinae is endemic to Africa and is thought to have diverged from other bovids during the early Miocene, around 20 million years ago. Impala are sexually dimorphic, as only males possess horns..

Cephalophinae consists of 18 species of duiker from 3 genera. Duikers are highly specialized and are resident to the tropical forests of Africa. All species are easily recognizable as they have the same basic body plan but differ significantly in size from one species to the next. Duikers are size-dimoprhic, however, unlike most bovids, females are slightly larger than males. Also unlike most other bovids, duikers are primarily frugivorous.

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Benefits

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Bovids, despite their important economic contributions to humans, can also have important detrimental effects. Zoonotic diseases transmitted by bovids to humans and domestic animals can have significant negative consequences, both physically and financially. For example, in less developed counties bovine tuberculosis can pose a significant economic threat for cattle farmers, and brucellosis, a bacterial disease that affects sheep, goats, cattle, elk, and deer, can be transmitted to humans by consuming undercooked contaminated meat and contaminated milk and dairy products. Bovine spongiform encephalopathy (BSE), more commonly known as Mad Cow Disease, is an infectious disease caused by an unknown agent, currently believed to be a modified protein. Cattle become infected when they are fed meat-and-bone meal that contains infected cattle by-products. Humans can contract BSE by consuming animal products from infected animals.

Bovids have been introduced world wide and in some locations have had severe detrimental impacts on the local environment. For example, goats were introduced by whalers to the Galapagos Island during the 18th century and have since caused extensive damage to the native ecosystem. In addition, introduced bovids compete with native animals for both food and habitat and can cause soil erosion due to overgrazing. Bovids, native and domestic, present a potential threat to various forms of agriculture by damaging and consuming crops.

Negative Impacts: injures humans (carries human disease); crop pest; causes or carries domestic animal disease

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
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Phil Myers, University of Michigan-Ann Arbor
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Benefits

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The domestication of artiodactyls for subsistence purposes lead to one of the most important cultural changes in human history, the transition from a purely hunter-gatherer society to a predominantly agricultural society. In the near east, around 10 thousand years ago (KYA), goat and sheep were domesticated purely for subsistence purposes, followed by the domestication of cows (7.5 KYA). Economically, cattle are the most important domesticated animal world wide. In 2001, the global population of domestic artiodactyls was greater than 4.1 billion, more than 31% of which consisted of cattle. In the United States, one of the worlds top 4 beef producers, beef production is the country's fourth largest industry, and in 2006, per capita beef consumption in the United States was nearly 66 pounds.

In addition to meat production, bovids are used for their milk, fur, skin, bone and feces. Goats and cattle are the primary producers of commercial milk and dairy products, sheep wool is used in the mass production of clothing, and manure is commonly used as fertilizer. For thousands of years humans have used bovids for hard labor tasks such as hauling materials, plowing fields, and transportation. Domestic bovids have also been used to control invasive plant species and enhance plant biodiversity through their selective feeding behavior.

Sport hunting of bovids generates millions of dollars annually. However, trophy hunting can alter the evolutionary dynamics of wild populations by imposing unnatural selective pressures for decreased ornamentation. Finally, bovids play an important role in the global ecotourism movement as various species are readily observable throughout much of their native habitat. Wildlife related tourism is especially popular in Eastern and Southern Africa and Central North America at various National Parks.

Positive Impacts: food ; body parts are source of valuable material; ecotourism ; research and education; produces fertilizer

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Associations

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As obligate herbivores, bovids can dramatically affect the abundance and diversity of plant communities. Predation, or the threat of predation, has been shown to decrease overgrazing by bovids. Bovids are host to a diverse array of endo- and ectoparasites. Many species of parasitic flatworms (Cestoda and Trematoda) and roundworms spend at least part of their lifecycle in the tissues of bovid hosts. Bovids are also vulnerable to various forms of parasitic arthropods including ticks, lice, mites (Psoroptes and Sarcoptes), keds, fleas, mosquitoes, and flies. Bovids also host various forms of parasitic protozoa, including trypanosomatids, coccidians, piroplasmids, and numerous species of Giardia. In addition, various forms of bacterial and viral pathogens play an important role in bovid health and population dynamics. For example, Brucella abortus, the bacteria that causes brucellosis, affects many bovid species and rhinderpest, also known as cattle plague, is a highly contagious viral disease caused by paramyxovirus that is especially prevalent in bovids. Unfortunately, evidence suggests that recent climate change is altering host-parasite dynamics across the globe, increasing transmission rates between populations of conspecifics and hybridization rates between host specific parasite forms.

Many bovids have mutualistic relationships with other animals. Cattle egrets and cowbirds regularly live amongst many bovid species, taking advantage of insects and parasites that feed on bovids, or feeding on insects and small animals that are forced out of hiding by movement and grazing. In addition to pest removal, mutualist species can alert them to the presence of predators. Bovids also create loosely formed interspecific groups with other large herbivores such as zebras, giraffes, and ostriches, which increases the chances for predator detection.

Although bovids can serve as host to numerous species of pathogenic bacteria and protozoa, in conjunction with anaerobic fungi, these organisms are one of the major reasons that bovids are as abundant and diverse as they are today. Bacteria help break down cellulose and comprise between 60 and 90% of the microbial community present in the gastrointestinal (GI) tract of bovids. Ciliated protozoa, which makes up 10 to 40% of the microbe community within the rumen, help bacteria break down cellulose, while also feeding on starches, proteins and bacteria. The presence of anaerobic fungi in the rumen has only been known since the early 1970's. These fungi make up between 5 to 10% of the rumen's microbial abundance and are thought to help break down the cell wall of ingested plant material. Bacteria and protozoa that pass from the upper to the lower regions of the GI tract represent a significant portion of the dietary nitrogen required by their host.

Ecosystem Impact: disperses seeds; soil aeration

Mutualist Species:

  • zebra, Equus
  • giraffe, Giraffa camelopardalis
  • ostrich, Struthio camelus
  • cattle egret, Bubulcus ibis
  • cowbird, Molothrus
  • rumen bacteria, Selenomonads
  • rumen bacteria, Oscillospira
  • rumen protozoa, Entodinium
  • rumen protozoa, Dasytricha
  • rumen protozoa, Diplodinia
  • rumen protozoa, Isotricha
  • rumen protozoa, Epidinia
  • rumen fungi, Neocallimastix
  • rumen fungi, Caecomyces
  • rumen fungi, Pyromyces
  • rumen fungi, Orpinomyces

Commensal/Parasitic Species:

  • nematodes, Nematoda
  • tapeworms, Cestoda
  • flukes, Trematoda
  • ticks, Ixodoidea
  • lice , Phthiraptera
  • flies, Diptera
  • mites, Psoroptes and Sarcoptes
  • keds, Hippoboscidae
  • fleas, Siphonaptera
  • mosquitoes, Culicidae
  • parasitic protozoa, Trypanosomatida
  • parasitic protozoa, Coccidia
  • parasitic protozoa, Piroplasmida
  • parasitic protozoa, Giardia
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bibliographic citation
Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Trophic Strategy

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Although bovids are obligate herbivores, they occasionally supplement their diet with animal products, and feeding strategies are correlated with body size. In general, small bovids are solitary specialized feeders that forage in dense, closed habitat, whereas large bovids tend to be gregarious and feed in open grassland habitats. As generalist herbivores, large bovids consume high-fiber vegetation, which contains more cellulose and lignin than the diet of forest dwelling species. However, because all bovids are obligate herbivores they support microbial communities within their rumen (bacteria, protozoa, and fungi), which help break down cellulose and lignin and converts high fiber forage into an abundant energy source.

In addition to the true stomach, or abomasum, all bovids have 3 additional chambers, or false stomachs, in which bacterial fermentation takes place. Bovids digest low-quality (i.e., low protein, high-fiber) food via four different pathways. First, gastric fermentation extracts lipids, proteins, and carbohydrates, which are then absorbed and distributed throughout the body via the intestines. Second, large undigested food particles form into a bolus, or ball of cud, which is regurgitated and re-chewed to help break down the cell wall of ingested plant material. Third, cellulose digestion via bacterial fermentation results in high nitrogen microbes that are occasionally flushed into the intestine, which are subsequently digested by their host. These high-nitrogen microbes serve as an important protein source for bovids. Finally, bovids can store large amounts of forage in their stomachs for later digestion. All bovids chew their cud, have four-chambered stomachs (1 true and 3 false stomachs) and support microorganisms that breakdown cellulose.

Each bovid subfamily has a unique feeding strategy. For example, members of Antilopinae are arid land gleaners and feed primarily on unevenly dispersed food resources. Bovinae species rely on both scattered and abundant forage and are fresh grass bulk grazers. Members of Caprinae are more generalized and flexible feeders and can often be found foraging in low-productivity habitats. Hippotraginae species are arid adapted grazers that generally rely on an unstable food supplies. Bovids from Reduncinae are valley grazers and depend on an abundant unstable food supply. Unlike most other bovids, members of Cephalophinae are primarily frugivorous and are known to follow canopy dwelling primates to collect dropped fruit.

Foraging Behavior: stores or caches food

Primary Diet: herbivore (Folivore )

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bibliographic citation
Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Distribution

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Although the greatest diversity of Bovidae occurs in Africa, bovids are also found throughout parts of Europe, Asia and North America. A number of bovid species, particularly those domesticated for subsistence, have been globally introduced, including Australia and South America.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Introduced )

Other Geographic Terms: holarctic ; cosmopolitan

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The Regents of the University of Michigan and its licensors
bibliographic citation
Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Habitat

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Bovids first evolved as grassland species, and most extant species are open grassland inhabitants. Bovid species richness is highest in the savannah of east Africa and the family has radiated to fill an enormous variety of ecological niches resulting in a wide range modifications to dental and limb morphology. For example, Bohor reedbuck and lechwe inhabit riparian and swampy landscape; springbok and oryx are found in deserts; bongo and anoa occupy dense forests; mountain goats and takin reside at high elevations; and musk ox are restricted to arctic tundra.

Numerous bovid species have been domesticated by humans. Goats and sheep were domesticated for subsistence purposes around 10 thousand years ago (KYA) in the near east, followed by the domestication of cattle around 7.5 KYA. While wild relatives of goats and sheep can still be found in their native habitat, the wild ancestors of domesticated cattle, aurochs, have been extinct in the wild for nearly 300 years. Currently, domesticated aurochs are kept on farms and as pets throughout parts of Eurasia.

Habitat Regions: temperate ; tropical ; polar ; terrestrial

Terrestrial Biomes: tundra ; taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp

Other Habitat Features: urban ; suburban ; agricultural ; riparian

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Life Expectancy

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Bovid lifespans are highly variable. Some domesticated species have an average lifespan of 10 years with males living up to 28 years and females living up to 22 years. For example, domesticated goats can live up to 17 years but have an average lifespan of 12 years. Most wild bovids live between 10 and 15 years, with larger species tending to live longer. For instance, American bison can live for up to 25 years and gaur up to 30 years. In polygynous species, males often have a shorter lifespan than females. This is likely due to male-male competition and the solitary nature of sexually-dimorphic males resulting in increased vulnerability to predation.

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Morphology

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Bovids display the characteristic long limbs and unique foot and unguligrade stance of artiodactyls. They are paraxonic, as the line of symmetry of the foot runs between the third and fourth digits. In most bovids, the lateral digits are either reduced or absent and the animal's weight is born on the remaining central digits. The third and forth metapodials are completely fused in bovids, resulting the cannon bone. The joint between the cannon bone and proximal phalanges includes four sesamoid bones that act as joint stops. The ulna and fibula is reduced and fused with the radius and tibia respectively. This arrangement provides for a wide angle of flexion and extension, but restricts lateral movement.

As a members of the suborder Ruminantia, bovids possess the trademark multi-chambered fore-gut adapted for cellulolytic fermentation and digestion. Thus, they are obligate herbivores, which is also reflected by their hypsodont and selenodont tooth morphology. Their upper incisors are absent and their upper canines are either reduced or absent. Instead of upper incisors, bovids have an area of tough, thickened tissue known as the dental pad, which provides a surface for gripping plant materials. The lower incisors project forward and are joined by modified canines that emulate the incisors. Their modified incisors are followed by a long toothless gap known as a diastema. Bovids have a generalized dental formula of I 0/3, C 0/1, P 2-3/3, M 3/3.

The distinguishing characteristic of the Bovidae family is their unbranching horns. The horns originate from a bony core known as the the cornual process (os cornu) of the frontal bone and are covered in a thick keratinized sheath. Horns are not shed like the antlers of cervids and most grow continuously. Except for Indian four-horned antelopes, horns occur in pairs and in a fascinating array of unique forms from curved daggers in mountain goat to the thick, rippled coils of greater kudu.

Bovids exhibit a wide range of sizes and pelage coloration and patterns. For example, gaurs have a maximum shoulder height of 3.3 m (10.82 ft) and a maximum weight of more than 1000 kg (2200 lbs), and pygmy antelope have a maximum shoulder height of 300 mm (1 ft) and a maximum weight of 3 kg (6.6 lbs). Forest and bush species tend to have shorter limbs and more developed hindquarters and cryptic pelage that helps them blend into their surroundings. Open habitat species have long, forelimbs that increase stride length and occasionally bold color patterns or stripes. These adaptations help bovids evade potential predators through the various mechanisms of hiding (cryptic coloration), escaping (increased stride length), or confusion (striped pelage).

Most bovids are sexually dimorphic. Males always have horns, which are used in ritualized fighting during the mating season. The horns of males tend to be more complex in design and more robust than those of females, which tend to be straighter, thinner, and simpler in design. Horns are present in females in approximately 75% of genera over 40 kg in mass and are usually absent in those less than 25 kg. This could be the result of differing life history strategies or the physiological cost of growing horns. Larger species are more likely to defend themselves against potential predators, and smaller species tend to retreat when threatened. In addition to sexual dimorphism in morphological characteristics, males also have better developed scent-glands than females, which are reduced or absent in species from the subfamily Bovinae.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: sexes alike; female larger; male larger; sexes shaped differently; ornamentation

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
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Tamatha A. Patterson, University of Michigan-Ann Arbor
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Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
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Phil Myers, University of Michigan-Ann Arbor
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Associations

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Bovids are an important food source for a variety of natural predators, and in Eastern and Southern Africa bovids are the primary food source for many pradator species including lions and cheetahs. On the African continent nearly all bovids are vulnerable to predation by lions and African wild dogs, but young, old and sick individuals are particularly susceptible. Leopards, spotted hyenas, cheetahs, Nile crocodiles, and side-striped jackals are also major predators of smaller bovid species. In North America, bovids are vulnerable to predation by grey wolves, brown bears, and cougar. Packs of wolves and adult bears are typically the only predators capable of taking down the largest bovids in North America, like American bison. On the continent of Asia, grey wolves and tigers, are predators of bovids. Leopards, dholes and mugger crocodiles are also capable of taking bovids as prey. There are some cases of Komodo dragons consuming goats and even water buffalo. Many predators like wild dogs and large cats are notorious for taking domesticated livestock, including domestic goats, domestic sheep, and cattle.

Bovids are formidable opponents and are capable of putting up an incredible fights against their predators. Strength in numbers, dangerous horns, powerful kicks, speed, and in some cases, sheer size are more than enough to deter most predation attempts. Muskox form tight knit circles of adults around their young, making an impenetrable wall against potential predators. Cape buffalo have been known to charge and kill lions. Many species of bovid are extremely fastest and use their speed to out maneuver predatory pursuers. Forest dwelling bovids, such as Bongo antelope have cryptic coats to help camouflage themselves in densely vegetated habitats.

Known Predators:

  • Lion Panthera leo
  • African wild dog Lycaon pictus
  • Leopard Panthera pardus
  • Spotted hyena Crocuta crocuta
  • Cheetah Acinonyx jubatus
  • Nile crocodile Crocodylus niloticus
  • Side-striped jackal Canis adustus
  • Grey wolf Canis lupus
  • Grizzly bear Ursus arctos horribilis
  • Brown bear Ursus arctos
  • Cougar Puma concolor
  • Tiger Panthera tigris
  • Dhole Cuon alpinus
  • Mugger crocodile Crocodylus palustris
  • Komodo Dragon Varanus komodoensis
  • Human Homo sapiens

Anti-predator Adaptations: cryptic

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Gomez, W.; T. Patterson; J. Swinton and J. Berini 2011. "Bovidae" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Bovidae.html
author
Whitney Gomez, University of Michigan-Ann Arbor
author
Tamatha A. Patterson, University of Michigan-Ann Arbor
author
Jonathon Swinton, University of Michigan-Ann Arbor
editor
John Berini, Special Projects
editor
Phil Myers, University of Michigan-Ann Arbor
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Diversity of Living Bovidae

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The family Bovidae (hollow-horned ruminants) is the largest and most diverse family in the mammal order Artiodactyla (even-toed ungulates). Bovids range in size from the 1.5 kg Royal Antelope (Neotragus pygmaeus) to the 1000 kg Common Eland (Taurotragus oryx) and Yak (Bos mutus). In most bovid species, females are hornless or have only small, fragile horns. Many of the more delicate-looking bovids are commonly known as "antelopes" (and some of the smaller antelopes are often known as "gazelles"). There are several bovids that are very familiar as domesticated animals, including cattle, buffaloes, sheep, and goats. Groves and Leslie (2011) provide an overview of the complex and tangled taxonomic history of Bovidae. With the availability of molecular phylogenetic data from DNA sequencing beginning in the 1990s, many aspects of bovid taxonomy andphylogeny have become clearer. A number of taxa formerly treated as single species are now recognized to include several distinct species. Groves and Leslie (2011) and Groves and Grubb (2011) recognize two subfamilies and 279 living bovid species:

Subfamily Bovinae. The subfamily Bovinae includes three tribes: Bovini, Boselaphini, and Tragelaphini. Members of this subfamily are medium to very large. Most are heavily built and adapted to living in heavy cover, often on soft (even swampy) terrain (only the American Bison [Bos bison] and Yak are found on open plains and in uplands).

Tribe Bovini (cattle and buffaloes). The 14 species in this tribe are placed in four genera: Bos, Bubalus, Syncerus, and Pseudoryx. A large wild species, the Aurochs (Bos primigenius) once ranged from Europe through southwestern Asia and North Africa. This species was domesticated (beginning, probably, in the Middle East or southwestern Europe), eventually yielding the domestic cattle we see today. As domestic cattle spread throughout the world, Aurochs were actively hunted by humans as competitors with domestic cattle and as dangerous game. The last known Aurochs was reportedly killed by a peasant in Poland in 1627. Efforts have been made to reconstruct cattle similar to the Aurochs through focused breeding efforts.

Cattle

Humped cattle, often known as Zebu, are descended from a wild species closely related to the Aurochs, Bos namadicus, that lived in India but is now extinct. Zebu have a floppy hump on the shoulder and a long, narrow dewlap and are well adapted to hot, dry conditions found in South Asia and drier parts of Southeastern Asia. African cattle, known as Sanga or Zeboid cattle, have small conical humps placed farther forward than those in Zebu and are apparently derived from an ancient cross between Zebu and humpless cattle.

The wild cattle of South and Southeast Asia are the Gaur (Bos gaurus), Kouprey (B. sauveli), and Banteng (Bos javanicus). The Gaur is among the largest of the bovids. Gaur populations in the border region of India, Bangladesh, Bhutan, and Burma have been domesticated, producing a smaller, stockier, short-horned form known as the "Mithan". Bantengs have been domesticated in central Indonesia, yielding a small domestic form known as Bali Cattle. The Kouprey, which was discovered by scientists only in the 1930s and inhabited the dry forests of northern Cambodia and adjacent parts of Thailand, Laos, and Vietnam, is tall and relatively slender with very large horns (especially in males) and a long, hanging dewlap. No Koupreys have been seen in the last several decades and the species may be extinct. Some genetic evidence suggests that Koupreys and Bantengs interbred in the past.

Bison

The two species commonly known as "bison" are not essentially distinct from other Bos (cattle) species, although they are sometimes segregated into a different genus, Bison. The American Bison has two forms. The first, the Plains Bison (Bos bison bison) is adapted to open, treeless plains and at one time millions of individuals roamed the North American plains. This number was reduced to around a thousand individuals by the 1880s as a result of large-scale slaughter. Populations have recovered somewhat thanks to concerted conservation efforts beginning early in the 20th century, although due to interbreeding with domestic cattle, many of the American Bison with us today are of mixed ancestry. A second form of American Bison, the Wood Bison (Bos bison athabascae) lived in the woodlands of northern Alberta and Mackenzie, Canada, until the early 20th century. In 1925, a large number of Plains Bison were released into the Wood Buffalo National Park, resulting in extensive interbreeding between the two forms, and it is now not clear just how distinct the Plains and Wood Bison were. The second bison species, the Wisent ("European Bison", Bos bonasus) was formerly found throughout the woodlands of central Europe, but was extinct in the wild by 1919. A third species, the Caucasian Bison (Bos caucasicus), persisted in a remnant population in the wild on the northwestern slopes of the Caucasus until 1927. As a consequence of concerted conservation efforts, the extinction of the European Bison was averted. However, of several hundred individuals in captivity today, around half are of pure lowland stock and half are the descendants of a single Caucasian bull crossed with lowland Wisents (European Bison reintroduced to the wild in Poland are all of pure lowland origin). The Yak of the Tibetan Plateau, which is adapted to living in open plains and uplands, is essentially a shaggy, long-horned bison.

Buffaloes

The buffaloes belong to two genera, Bubalus (Asian buffaloes) and Syncerus (African buffaloes). The Asian Wild Buffalo (Bubalus arnee) lineage gave rise to the two forms of domestic water buffalo, the "swamp buffalo" of Southeast Asia and the "river buffalo" of South Asia (and widely introduced elsewhere). Several dwarf Bubalus species have evolved on islands. Although Syncerus was long treated as including just a single geographically variable species, new data have led to the recognition of several distinct Syncerus species.

Saola

Pseudoryx includes just a single species, the Saola (P. nghetinhensis), found in the mountains of Laos and Vietnam and know to science only since the early 1990s.

Tribe Boselaphini (Nilgai and Chowsingha). The 2 species in this tribe are both found in India. Although they share many anatomical characters, the Nilgai (Boselaphus tragocamelus) is around ten times the size of the Chowsingha (Tetracerus quadricornis). The male Chowsingha is the only wild mammal with four horns (the females lack horns altogether and in some populations males have only two horns). The Nilgai, Chowsingha, and Saolo are the only Bovinae with facial glands (located in front of the eyes).

Tribe Tragelaphini (bushbucks, kudus, and elands). The 24 species in this tribe are placed in five genera: Nyala (the Nyala [N. angasii]), Tragelaphus (Bongo [T. eurycerus], Gedemsa [T. buxtoni], 8 bushbuck species, 5 sitatungas), Ammelaphus (2 lesser kudu species), Strepsiceros (4 greater kudu species), and Taurotragus (2 eland species, the Giant Eland and Common Eland; the Giant Eland is named for its enormous horns and actually has a smaller body than the Common Eland; both sexes of elands have horns). Genetic and chromosomal studies have clarified a number of questions re. species boundaries and relationships within this tribe, but also suggest a complex ancient history of genetic introgression among some lineages. The horns of Tragelaphini are keeled, with the keel starting above the eye and following the spiral of the horn nearly to its tip. The elands are the tallest of the African antelopes, followed by the greater kudus; male greater kudus have the longest and most widely spiraled horns of all African antelopes.

Subfamily Antilopinae. The Subfamily Antilopinae includes nine tribes: Neotragini, Aepycerotini, Antilopini, Reduncini, Hippotragini, Alcelaphini, Caprini, Cephalophini, and Oreotragini. This subfamily includes species ranging from the smallest bovid to medium- and large-sized forms. Many species have brownish upperparts and lighter (often strikingly white) underparts, frequently with a dark band along the flanks separating the two zones. The face often has longitudinal dark and light stripes from the eye to the nostril, sometimes reduced to a white eye ring. The horns are transversely ringed, sometimes strongly so. There are nearly always prominent glands on the face, in front of the eyes, and in the feet (at least the forefeet). With the exception of mountain-dwelling species, members of the Antilopinae tend to be more lightly built than bovines and most live in relatively open country (the tribes Cephalophini and Neotragini being exceptions). Antilopine social structure usually involves territorial males and separate herds of females.

Tribe Neotragini (dwarf antelopes). Although other small antelopes used to be included in this tribe, it is now clear that only the 5 species in the genus Neotragus actually belong in th Neotragini. Neotragus includes the tiny Royal Antelope of West Africa, the Dwarf Antelope (N. batesi) of Central Africa, and the three suni species of East Africa. The horns of these antelopes (which are present only in males) are short and grow backward approximately in the plane of the forehead. Neotragus antelopes are the only African bovids other than Cephalophini (duikers), Forest Buffalo (Syncerus nanus), and Bongo that are associated with closed forest.

Tribe Aepycerotini. This tribe includes only the 2 species of impalas, the Black-faced Impala (Aepyceros petersi) and Common Impala (A. melampus). The very slender horns have strong transverse ridges and a single wide twist, with tips that turn upward. Impalas have inguinalglands but no facial glands, carpal glands, or foot glands. Impalas are found in eastern and southern Africa.

Tribe Antilopini (gazelles, dik-diks, and relatives). The 64 species in this tribe are placed in 13 genera: Raphicerus, Antidorcas, Ammodorcas, Litocranius, Saiga, Antilope, Nanger, Gazella, Eudorcas, Dorcatragus, Madoqua, Ourebia, and Procapra. Members of this tribe are all found in open grassland or dry woodlands. All Antilopini have a slender build and most have disruptive coloration, typically sandy toned or reddish or gray above and white below, with the upper and lower zones often separated by a dark band along the lower flanks. The buttocks are white, with this area demarcated by another dark stripe on either side. A white eye ring on the face often continues downward to the nostrils as a white stripe, flanked above by a dark stripe. Large preorbital glands are present. In most species in this tribe, only males have horns (however, in most of the species commonly known as gazelles, the females have small, slender horns). The horns in larger species have transverse ridges, at least on the front surface; in smaller species, the horns are more or less smooth. Five major clades are recognized within this tribe:

1) 4 spp. The genus Raphicerus includes the Steenbok (R. campestris) and 3 species of grysboks from southern and eastern Africa. These small antelopes, which have straight, slender, vertically oriented horns are found in brushland, high-grass country, and woodland.

2) 40 spp. This clade includes small and medium-sized antelopes of arid and semi-arid country: the springboks (Antidorcas, 3 species), gerenuks (Litocranius, 2 spp.), and the Dibatag (Ammodorcus clarkei) of Africa; the Blackbuck (Antilope cervicapra) of the South Asian plains, in which males are black and females are tawny; and the gazelles. All of these have facial glands, foot glands, and carpal glands, and most have inguinal glands. At one time, the Blackbuck was found throughout the open plains of India and Pakistan, but the population of at least a million in the 19th century has dropped to a few tens of thousands. The three genera known as gazelles (which are not actually a phylogenetically coherent group) are placed in the genera Gazella (21 species of small gazelles of desert country from the Sahara to the Gobi); Eudorcas (larger gazelles, including 3 species of red-fronted gazelles and 2 species of Thompson's gazelles from the Sahel and East African savanna); and Nanger (5 spp., including the large Grant's Gazelle [N. granti], Soemmerring's Gazelle [N. soemmerringii], and Dama Gazelle [N. dama]). The unusual-looking Saiga (Western Saiga [S. tatarica] and Mongolian Saiga [S. mongolica]) of the Central Asian steppes also fall within this clade.

3) 13 spp. This clade includes the dik-diks (Madoqua, 12 spp.), which live in arid thornbush country in the same areas as the gerenuks and the Beira (Dorcatragus megalotis). The horns (present only in males) are tiny. Dik-diks have facial glands, but the Beira does not. Dik-dik's are well adapted for hot, dry environments.Guenther's Dik-diks (M. guentheri), for example, pant rather than sweat and their blood is cooled by an enormous nasal cavity, the flexible proboscis acting like a bellows to increase airflow. These dik-diks licks droplets that condense on the outside of the nose and their kidneys are reportedly second only to those of jerboas and kangaroo rats (desert-adapted rodents) in their ability to concentrate urine.

4) 4 spp. The Oribi (Ourebia, 4 spp.) are small to medium-sized antelopes living in grassland and bush country throughout sub-Saharan Africa. Oribi have facial glands and large glandular carpal tufts and a bare patch of skin beneath each ear (as in reedbucks, to which they are not especially closely related). Oribi have long legs, a fairly long neck, and large ears. The horns, which are thin spikes, are present only in males.

5) 3 spp. Procapra includes the Mongolian Gazelle (P. gutturosa), Przewalski's Gazelle (P. przewalskii), and Tibetan Gazelle (P. picticaudata). Members of this group have noticeably short tails lack carpal glands, have only small facial glands, if any, and lack the striking facial patterns of "true" gazelles. Females lack horns.

Tribe Reduncini. The 22 species in this tribe include the reedbucks, waterbucks, and relatives. Most are water-dependent medium to large antelopes. Horns, which are transversely ridged, are present only in males. The three genera in this tribe are Redunca (9 spp.), Kobus (8 spp.), and Pelea. Redunca (reedbuck) are found in open habitats of sub-Saharan Africa, mainly in moist environments. Kobus includes three species-groups (sometimes treated as distinct genera). The first of these, the waterbucks, are found throughout the plains of sub-Saharan Africa, always near water. The rather small kobs have large inguinal and preorbital glands and somewhat S-shaped horns and live in well-watered plains from West Africa to the Sudan and Uganda. The Puku (K. vardoni) lives much farther south, from Botswana to Mozambique. The lechwe, which are intermediate in size between waterbucks and kobs, lack preorbital glands and have long, sweeping S-shaped horns and reddish or blackish shaggy coats. Lechwe are among the most strongly water-associated antelopes, second only to most of the sitatunga, and are found in the floodplains of Zambia and neighboring countries and in Sudan. The Rhebok (Pelea capreolus), found in the highveld of South Africa, is the smallest member of this tribe. Rheboks have glands in the feet (but no other skin glands), soft wooly gray fur, and short, straight, upright horns.

Tribe Hippotragini. The 10 species in this tribe are placed in 3 genera: Hippotragus, Oryx, and Addax. Members of this tribe are medium-sized to large antelopes with long horns in both sexes. The horns are parallel or only slightly divergent. They curve back or are nearly straight or spiraled and are clearly ringed all along their length. Hippotragus includes 3 species found in well-watered high-grass habitats: Southern Sable Antelope (H. niger) and Roosevelt's Sable Antelope (H. roosevelti), found in southern and eastern Africa, and the Roan Antelope (H. equinus), which occurs throughout the grasslands of sub-Saharan Africa, An additional species, the Blaubok (H. leucophaeus) lived in the Cape provinces of South Africa but was wiped out by Boer expansion. When first discovered, it was known only from a small area of the southwestern Cape. That population was extinct by 1800, but another population may have persisted in the Free State province until the 1850s. Oryx includes desert and semi-desert species in both Africa and Arabia. Most species have very long, almost straight, backward-pointing horns. The Beisa Oryx of East Africa provides an example, not uncommon among bovids, of a group long treated as a variable single species turning out to represent several distinct species, in this case Beisa Oryx (Oryx beisa), Galla Oryx (O. gallarum), and Fringe-eared Oryx (O. callotis). Addax includes just a single species, the Addax (A. nasomaculatus) of the Sahara. The Addax resembles an Oryx but with spiral horns, large spreading hooves, and a bushy tuft of hair on the forehead.

Tribe Alcelaphini. Members of this tribe are large antelopes with horns in both sexes. There are 23 species in this tribe, placed in four genera: Beatragus (1 species), Alcelaphus (hartebeest, 9 spp.), Damaliscus (11 spp.), and Connochaetes (wildebeest, 5 spp.). Hartebeest have strikingly long faces and horns that are raised on a high shelf-like pedicle. They are found in grasslands across sub-Saharan Africa and at one time their range extended into North Africa and even Israel, but this northernmost representative, the Bubal Hartebeest (A. buselaphus) ,went extinct in the 1920s. Hartebeest horns are always much thicker and sturdier in males than in females. Male hartebeest fight by dropping to their knees and battering each other and the positioning of their horns on raised pedicles displaces the impact of the blows away from the braincase. Damaliscus species have elongated faces, like hartebeests, but do not have elongated horn pedicles. The Bontebok (D. pygargus) was driven nearly to extinction by the early Dutch settlers of South Africa, but today is no longer in danger of extinction. Most of the other Damaliscus species are fairly numerous, but the Uasin Gishu Topi (D. selousi), of western Kenya, has not been recorded in recent years and is likely extinct.The Critically Endangered Herola (Beatragus hunteri) is known in historical times only from the region between the Tana River in Kenya and the Juba River in Somalia. The best known members of this tribe are the wildebeests. Wildebeest are found only in southern and eastern Africa, not reaching Sudan or West Africa.

Tribe Caprini. The 61 species in this tribe include the sheep, goats, and relatives. Most species live in hilly or mountainous regions and tend to have short, sturdy metapodials (shin bones). In nearly all species, both sexes have horns.

1. The Chiru (Pantholops hodgsonii) is found on the Tibetan Plateau. It has very long upright horns (present in males only) and has inguinal glands but no glands on the face or feet. Although its numbers are much reduced from the past, it still occurs in herds of thousands. Its wool is extemely fine and much sought after.

2. Oreamnos includes just a single species, the Rocky Mountain Goat (O. americanus) of the higher mountains of western North America. It is a stocky, shaggy, white goat-like animal with short, simple back-curved horns

3. Budorcas includes 4 species known as takins and found on the high mountain slopes of western and central China and the eastern Himalayas. They apparently have no glands on the face or in the feet or groin.

4. Rupicapra includes 6 species known as chamois. These occur in the high mountains of Europe from Spain east to the Caucasus. Chamois have gazelle-like facial stripes, a lighter underside, and slender horns that rise vertically and hook backward at the tips. They have small foot glands and no glands on the face, but have a pair of glands behind the horns.

5. Nemorhaedus, Capricornis, and Ovibos form a clade together. Nemorhaedus is composed of six species of gorals, small, compactly built "goat-antelopes" with short back-curved horns. Gorals live in high, steep areas from northern Thailand west through the Himalayas to Kashmir and north to the Russian Far East. The 7 species of serows (Capricornis) look superficially like very large gorals, with the same short, back-curved horns, but unlike Gorals, they have facial glands and a very short tail. Serows live at relatively low elevations, but also in steep country as far south as Sumatra and north to central China and in the Himalayas. Ovibos includes a single species, the Muskox (Ovibos moschatus), which is a heavily built animal of the North American Arctic with very long shaggy hair and white fibrous horns that curve outward, down, then up. Like the related serows, Muskoxen have very short tails, but they have only small facial glands.

6. The goats and relatives have been the focus of much taxonomic debate. Molecular data have now confirmed that blue sheep, the Aoudad (Ammotragus lervia), and the tahrs are more closely related to goats than to sheep. The Greater Blue Sheep (Pseudois nayaur) and Lesser Blue Sheep (P. schaeferi) are superficially sheep-like, but have no facial glands and the foot glands are at most rudimentary. The Greater Blue Sheep is abundant in the higher alpine zone of the Himalayas and the eastern margins of the Tibetan Plateau and the Dwarf Blue Sheep occurs in the arid upper reaches of the Yangtze Gorge, separated from the alpine zone by a 1000 m wide forest belt. The Aoudad, also known as the "Barbary Sheep", is found in the semi-arid massifs of the Sahara. It is one of only three species in the tribe Caprini occurring in Africa (the other two being Nubian Ibex [Capra nubiana] and Walia Ibex [C. walie]). Like blue sheep, Aoudads have no glands in the face or feet. The Himalayan Tahr (Hemitragus jemlahicus) is found on the southern slopes of the Himalayas in very steep, rocky areas. Recent work has indicated that what has been treated as a three closely related Hemitragus species is actually several species: H. jemlahicus, related to goats (Capra),b ut even more closely related to Pseudois; Nilgiri Tahr (Nilgiritragus hylocrius), closely related to sheep (Ovis); and Arabian Tahr (Arabitragus), the sister group to Ammotragus. The Arabian Tahr lives in the mountains of northeastern Oman and the far north of the United Arab Emirates. The genus Capra includes the goats. All Capra have beards and fairly short, triangular tails, bare underneath and with strong-smelling glands at the base. There are no facial glands and if foot glands are present they are only found on the front feet. Capra includes the ibexes, most of which live in high mountains from the Altai in Siberia west to Spain and south to Ethiopia, although the Nubian Ibex lives in lower but steep hills in Israel and along both sides of the Red Sea. The Wild Goat (Capra aegagrus), or "Bezoar", is the only wild representative of the "true goats". It lives in steep hilly areas from Turkey east to Pakistan and is probably the sole ancestor of domestic goats, although some authorities have suggested that a few breeds of goats may be partly derived from the Markhor (Capra falconeri), which lives in hilly areas of Pakistan, Indian Kashmir, Afganistamn, Tajikstan, and Uzbekistan. In Pakistan, Markhors and Wild Goats have interbred to produce a hybrid with loosely spiraled horns known as the "Chiltan Goat". Molecular phylogenetic studies suggest a complex history of ancient hybridization between lineages within the Capra clade.

7. The genera Ovis (sheep) and Nilgitragus form a clade. Sheep have both facial and foot glands (whether the Nilgiri Tahr has facial glands is not yet known). The Nilgiri Tahr lives in the alpine zones of the Western Ghats of India. Wild sheep (Ovis) are found from Turkey through West and Central Asia into northeastern Siberia, Alaska and the Rocky Mountains and desert hills of western North America. The American species inhabit steep terrain, almost like goats. Domestic sheep are derived from members of the Ovis gmelini group. Most lines of domestic shhep have been bred to increase the production of winter underwool at the expense of the normal contour hairs. Domestic breeds not selected for their wool often look very much like their wild ancestors (e.g., the feral "Mouflons" from the Mediterranean islands of Corsica, Sardinia, and Cyprus).

Tribe Cephalophini. This tribe includes the many small forest-dwelling antelopes known as duikers. There are 41 species, placed in three genera: Philantomba, are very small and are found throughout the forest zones of Africa, from the main rainforest belt of West and Central Africa through the highland and coastal forests of eastern and southern Africa. One species (Verheyen's Duiker [P. walteri], from West Africa) was described only in 2010. Duikers seem to be very abundant in rainforests. They are often a mainstay of the bushmeat trade yet appear nevertheless to remain common. Sylvicapra males have a long, narrow tuft of hair between the horns. Two Sylvicapra species occur in western Africa and a third species is found throughout non-forested regions in sub-Saharan Africa. Cephalophus includes more heavily built duikers. Two West African species (Banded Duiker [C. zebra] and Jentink's Duiker [C. jentinki]) and one East African species (Ader's Duiker [C. adersi]) are Critically Endangered.

Tribe Oreotragini. Oreotragus (the klipspringers) includes 11 species of small, stocky antelopes with short, thick hooves. They are found on isolated rocky outcrops so their distribution is patchy.

(Groves and Grubb 2011 and references therein; Groves and Leslie 2011 and references therein)

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Bovidae

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The Bovidae are the biological family of cloven-hoofed, ruminant mammals that includes bison, African buffalo, water buffalo, antelopes, sheep, goats, muskoxen, and domestic cattle. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the family Bovidae consists of eight major subfamilies apart from the disputed Peleinae and Pantholopinae. The family evolved 20 million years ago, in the early Miocene.

The bovids show great variation in size and pelage colouration. Excepting 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 diurnal. Social activity and feeding usually peak during dawn and dusk. Bovids typically rest before dawn, during midday, and after dark. They have various methods of social organisation and social behaviour, which are classified into solitary and gregarious behaviour. Bovids use different forms of vocal, olfactory, and tangible communication. Most species alternately feed and ruminate throughout the day. While small bovids forage in dense and closed habitat, larger species feed on high-fiber vegetation in open grasslands. Most bovids are polygynous. Mature bovids mate at least once a year and smaller species may even mate twice. In some species, neonate bovids remain hidden for a week to two months, regularly nursed by their mothers; in other species, neonates are followers, accompanying their dams, rather than tending to remain hidden.

The greatest diversities of bovids occur in Africa. The maximum concentration of species is in the savannas of eastern Africa. Other bovid species also occur in Europe, Asia, and North America. Bovidae includes three of the five domesticated mammals whose use has spread outside their original ranges, namely cattle, sheep, and goats. Dairy products such as milk, butter, and cheese are manufactured largely from domestic cattle. Bovids also provide leather, meat, and wool.

Etymology

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 family Bovidae is 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 known extinct species.[3]

Molecular studies have supported monophyly in the family Bovidae (a group of organisms comprises an ancestral species and all their descendants).[4][5] The number of subfamilies in Bovidae is disputed, with suggestions of as many as ten and as few as two subfamilies.[5] However, molecular, morphological and fossil evidence indicates the existence of eight distinct subfamilies : Aepycerotinae (consisting of just 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, three extinct subfamilies are known: Hypsodontinae (mid-Miocene), Oiocerinae (Turolian) and the subfamily Tethytraginae, which contains Tethytragus (mid-Miocene).[6][7]

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.[8]

A controversy exists 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,[9] though the grey rhebok, the sole species of Pelea, is highly different from kobs and reduncines in morphology.[10] Pantholops, earlier classified in the Antilopinae, was later placed in its own subfamily, Pantholopinae. However, molecular and morphological analysis supports the inclusion of Pantholops in Caprinae.[11]

Below is a cladogram based on Gatesy et al. (1997) and Gentry et al. (1997)

Bovidae Boodontia (Bovinae)  

Tragelaphini (kudus, nyalas etc.)"The

   

Bovini (bison, buffalo, cattle, etc.)"Birds

    Aegodontia Antilopinae

Antilopini (gazelles, springbok etc.)"The

   

Neotragini (dik-diks etc.)"The

       

Cephalophinae (duikers etc.)"The

   

Reduncinae (kobs, reedbucks, waterbucks etc.)"The

     

Aepycerotinae (impala)"The

    Caprinae

Ovibovini (takin, muskox)

   

Caprini (chamois, sheep, ibexes, goats etc.)"Walia

     

Hippotraginae (sable antelopes, oryxes etc.)"The

   

Alcelaphinae (hartebeest, topi, wildebeest etc.)"The

       

Evolution

Early Miocene and before

"
Skull of Eotragus sansaniensis, a species of the ancient bovid genus Eotragus

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.[12] Eotragus, the earliest known bovid, weighed 18 kg (40 lb) and was nearly the same in size as the Thompson's gazelle.[13] Early in their evolutionary history, the bovids split into two main clades: Boodontia (of Eurasian origin) and Aegodontia (of African origin). This early split between Boodontia and Aegodontia 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 territory of the other.[14] The tribes Bovini and Tragelaphini diverged in the early Miocene.[15] Bovids are known to have reached the Americas in the Pleistocene by crossing the Bering land bridge.[13]

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 became extinct in the middle Pleistocene.[5] 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.[15] The Bovinae are believed to have diverged from the rest of the Bovidae in the early Miocene.[16] The Boselaphini became extinct in Africa in the early Pliocene; their latest fossils were excavated in Langebaanweg (South Africa) and Lothagam (Kenya).[17]

Middle Miocene

The middle Miocene marked the spread of the bovids into China and the Indian subcontinent.[13] According to Vrba, the radiation of the subfamily Alcelaphinae began in the latter part of middle Miocene.[5] The Caprinae tribes probably diverged in the early middle Miocene. The Caprini emerged in the middle Miocene, and seem to have been replaced by other bovids and cervids in Eurasia.[18] 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.[15]

Late Miocene

By the late Miocene, around 10 Mya, the bovids rapidly diversified, leading to the creation of 70 new genera.[13] This late Miocene radiation was partly because many bovids became adapted to more open, grassland habitats.[12] The Aepycerotinae first appeared in the late Miocene, and no significant difference in the sizes of the primitive and modern impala has been noted.[19] Fossils of ovibovines, a tribe of Caprinae, in Africa date back to the late Miocene.[15] The earliest Hippotragine fossils date back to the late Miocene, and were excavated from sites such as Lothagam and Awash Valley.[15] The first African fossils of Reduncinae date back to 6-7 Mya.[20] Reduncinae and Peleinae probably diverged in the mid-Miocene.[5]

Characteristics

"
Bovids have unbranched horns.

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.[21] 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 as much as 1,000 kg (2,200 lb) and stands 2–3 m (6.6–9.8 ft) high at the shoulder.[22] 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) tall.[23] The royal antelope, in sharp contrast, is only 25 cm (9.8 in) tall and weighs at most 3 kg (6.6 lb).[24] The klipspringer, another small antelope, stands 45–60 cm (18–24 in) at the shoulder and weighs just 10–20 kg (22–44 lb).[25]

Differences occur in pelage colouration, ranging from a pale white (as in the Arabian oryx)[26] to black (as in the black wildebeest).[27] However, only the intermediate shades, such as brown and reddish brown (as in the reedbuck), are commonly observed.[28] In several species, females and juveniles exhibit a light-coloured coat, while those of males darken 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 by the season.[29] Scent glands and sebaceous glands are often present.[21]

"
The gemsbok has conspicuous markings on its face, which conceal the eye, and on its legs. These may have a role in communication.[30]

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.[31] Many species, such as gazelles, may be made to look flat, and hence to blend into the background, by countershading.[32] The outlines of many bovids are broken up with bold disruptive colouration, the strongly contrasting patterns helping to delay recognition by predators.[33] 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.[30]

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 four-horned antelope[34] and the Jacob sheep.[35][36] The unique horn structure is the only unambiguous morphological feature of bovids that distinguishes them from other pecorans.[37][38] A high correlation exists 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.[39] Males with horns directed inwards are monogamous and solitary, while those with horns directed outwards tend to be polygynous. These results were independent of body size.[40]

Male horn development has been linked to sexual selection,[41][42] 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.[28] However, the presence of horns in females is likely due to natural selection.[41][43] The horns of females are usually smaller than those of males, and are sometimes of a different shape. The horns of female bovids are believed to have evolved for defence against predators or to express territoriality, as nonterritorial females, which are able to use crypsis for predator defence, often do not have horns.[43] Females possess horns only in half of the bovid genera, and females in these genera are heavier than those in the rest. Females use horns mainly for stabbing.[44]

"
American bison skeleton (Museum of Osteology)

Anatomy

"
Dental pad of a domestic bovid: Note the absence of upper incisors and canines and the outward projection of the lower teeth.

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 fluctuates through the day; for instance, in goats the temperature can change slightly from nearly 37 °C (99 °F) in the early morning to 40 °C (104 °F) 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.[3][21]

Dentition

Most bovids bear 30 to 32 teeth.[28] 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.[45] The general dental formula for bovids is 0.0.2-3.33.1.3.3. Most members of the family are herbivorous, but 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.[3]

Ecology and behaviour

"
Blackbuck antelopes

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.[28] Small bovids such as the klipspringer, oribi, and steenbok are generally solitary and territorial. They hold small territories into which other members of the species 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.[46] The offspring disperse at the time of adolescence, and males need must acquire territories prior to mating.[3] The bushbuck is the only bovid that is both solitary and not 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.[47]

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 must 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.[28] 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 they may acquire his territory.[48] Lek mating, where males gather together and competitively display to potential mates, is known to exist among topis, kobs, and lechwes.[49] The tragelaphines, cattle, sheep, and goats are gregarious and not territorial. In these species, males must gain absolute dominance over all other males, and fights are not confined to territories. Males, therefore, spend years in body growth.[28]

Activity

"
Blue wildebeest fighting for dominance

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 flehmen response. Bovids usually stand motionless, with the head high and an intent stare, when they sense danger. Some like the impala, kudu, and eland can even leap to heights of a few feet.[28] Bovids may roar or grunt to caution others and warn off predators.[3] Bovids such as gazelles stot or pronk in response to predators, making high leaps on stiff legs, indicating honestly both that the predator has been seen, and that the stotting individual is strong and not worth chasing.[50]

"
Stotting or pronking by a young springbok signals honestly to predators such as cheetahs that it is a fit and fast individual, not worth chasing.

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.[51][52]

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.[28][51][53]

Diet

"
Bovids are herbivores, feeding on grass, foliage, and plant products.

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.[28] 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 Sylvicapra) primarily consume fruits.[3] Reduncinae and Hippotraginae species depend on unstable food sources, but the latter are specially adapted to arid areas. Members of Caprinae, being flexible feeders, forage even in areas with low productivity. Tribes Alcelaphini, Hippotragini, and Reduncini have high proportions of monocots in their diets. On the contrary, Tragelaphini and Neotragini (with the exception of Ourebia) feed extensively on dicots.[54] No conspicuous relationship exists between body size and consumption of monocots.[55]

Sexuality and reproduction

"
Juvenile sheep (lamb) near its mother

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.[56][57] 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.[58] On the contrary barbary sheep females may give birth to offspring even before they have gained sexual maturity.[59] 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.[27]

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 photoperiod plays a pivotal role. Other factors that have a significant influence on this cycle include the temperature of the surroundings, nutritional status, social interactions, the date of parturition and the lactation period. A study of this phenomenon concluded that goats and sheep are short-day breeders. Mating in most sheep breeds begins in summer or early autumn.[60] Mating in sheep is also affected by melatonin, that advances the onset of the breeding season;[61] and thyroxine, that terminates the breeding season.[62] 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.[28] 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.[28][56]

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.[58] 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).[56][57]

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).[63][64] 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.[65]

Distribution

"
Eland occur in grasslands of Africa.

Most of the diverse bovid species occur in Africa. The maximum concentration is in the savannas of eastern Africa. Depending on their feeding habits, several species have radiated over large stretches of land, and hence several variations in dental and limb morphology are observed. Duikers inhabit the equatorial rainforests, sitatunga, and lechwe occur near swamps, eland inhabit grasslands, springbok and oryx occur in deserts, bongo and anoa live in dense forests, and mountain goats and takin live at high altitudes.[28] A few bovid species also occur in Europe, Asia, and North America. Sheep and goats are found primarily in Eurasia, though the Barbary sheep and the ibex form part of the African fauna. The muskox is confined to 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][56]

Interaction with humans

Domesticated animals

"
Zebu oxen in Mumbai

The domestication of bovids has contributed to shifting the dependence of human beings from hunting and gathering to agriculture. The Bovidae includes three of the five large domesticated herbivores whose use has spread outside their original ranges, namely cattle, sheep, and goats; all are from Eurasia, and are now found across the world. The other two species are the horse and pig. Other large bovids that have been domesticated but which remain within the ranges of their wild ancestors are the water buffalo (from the Indian water buffalo), domestic yak (from the wild yak), zebu (from the Indian aurochs), gayal (from the gaur) and Bali cattle (from the banteng).[56] Some antelopes have been domesticated including the scimitar oryx, 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.[66]

Animal products

"
Merino wool is the most valued, with great fineness and softness.

Dairy products such as milk, butter, ghee, 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,[67] while sheep milk is used to make blue Roquefort cheese in France.[68] Beef is an excellent source of zinc, selenium, phosphorus, iron, and B vitamins.[69] Bison meat is lower in fat and cholesterol than beef, but has a higher protein content.[70]

Bovid leather is tough and durable, with the additional advantage that it can be made into leathers of varying thicknesses - from soft clothing leather to hard shoe leather. While goat and cattle leather have a wide variety of use, sheepskin is suited only for clothing purposes.[71] Wool from Merino hoggets is the finest and most valuable. 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[72] and carpets.

"
Drinking horn made by Brynjólfur Jónsson of Skarð, Iceland, 1598

Bone meal is an important fertilizer rich in calcium, phosphorus, and nitrogen, effective in removing soil acidity.[73] Bovid horns have been used as drinking vessels since antiquity.[74]

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.[75] 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.[76] The sheep, synonymous with the goat in Chinese mythology, is the eighth animal of the Chinese zodiac, and a symbol of filial piety.[77]

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Bovidae: Brief Summary

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The Bovidae are the biological family of cloven-hoofed, ruminant mammals that includes bison, African buffalo, water buffalo, antelopes, sheep, goats, muskoxen, and domestic cattle. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the family Bovidae consists of eight major subfamilies apart from the disputed Peleinae and Pantholopinae. The family evolved 20 million years ago, in the early Miocene.

The bovids show great variation in size and pelage colouration. Excepting 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 diurnal. Social activity and feeding usually peak during dawn and dusk. Bovids typically rest before dawn, during midday, and after dark. They have various methods of social organisation and social behaviour, which are classified into solitary and gregarious behaviour. Bovids use different forms of vocal, olfactory, and tangible communication. Most species alternately feed and ruminate throughout the day. While small bovids forage in dense and closed habitat, larger species feed on high-fiber vegetation in open grasslands. Most bovids are polygynous. Mature bovids mate at least once a year and smaller species may even mate twice. In some species, neonate bovids remain hidden for a week to two months, regularly nursed by their mothers; in other species, neonates are followers, accompanying their dams, rather than tending to remain hidden.

The greatest diversities of bovids occur in Africa. The maximum concentration of species is in the savannas of eastern Africa. Other bovid species also occur in Europe, Asia, and North America. Bovidae includes three of the five domesticated mammals whose use has spread outside their original ranges, namely cattle, sheep, and goats. Dairy products such as milk, butter, and cheese are manufactured largely from domestic cattle. Bovids also provide leather, meat, and wool.

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