Tragelaphini Temporal range: Middle Miocene – present, [1] | |
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Giant eland showing the characteristic spiral-horns seen in this group of bovines. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Family: | Bovidae |
Subfamily: | Bovinae |
Tribe: | Tragelaphini Blyth, 1863 sensu Sokolov, 1953 |
Type species | |
Antilope sylvatica Pallas, 1766 | |
Genera | |
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Synonyms | |
The tribe Tragelaphini (sometimes referred to by some authors as "Strepsicerotini" [2] [3] ), or the spiral-horned antelopes, are bovines that are endemic to sub-Saharan Africa. These include the bushbucks, kudus, and the elands. The scientific name is in reference to the mythical creature the tragelaph, a Chimera with the body of a stag and the head of a goat. They are medium-to-large, tall, long-legged antelopes characterized by their iconic twisted horns and striking pelage coloration patterns (most common is the distinctive white, vertical barring). [4]
Despite being among the largest species of antelope, they are actually more closely related to cattle (Bos taurus), and together along with a few apparent Asiatic species belong to the subfamily Bovinae. [4] [5] [6] While the group's evolutionary history occurred in Africa, there have been fossil species that have been found in Eurasia (which may also be the place of origin for this group). [7] [8] The number of genera and species is debated as some consider there to be one or two genera with nine species, while others consider there to be five genera and 25 species. [6] In general, spiral-horned antelopes can be roughly divided into two groups: robust forms (which only consists of the two eland species, Taurotragus ) and gracile forms (the rest of them, in the genus Tragelaphus , although this taxon is an unnatural grouping, and might warrant additional genera). [5] [6] [1]
Spiral-horned antelopes are browsers, found in a wide variety of environments both arid and humid, including semi-deserts, savannas, rainforests and mountains. [1] [4] In all these environments, however, they prefer to live in dense bush or thicket, which offer concealment from their natural predators. Considered among the most beautiful and charismatic bovids, the various species of spiral-horned antelopes are popular in zoos and game reserves. [9] The two eland species have been ranched as alternatives to cattle, being hardy in extreme environments, relatively placid in character, and nutritionally superior as a meat source. [10] [11] [12] [13]
The tribe name "Tragelaphini" was published as a subfamily ("Tragelaphinae") by British zoologist Edward Blyth in 1863, and was downgraded to tribe by Russian zoologist Vladimir Sokolov in 1953. [2] [3] It refers to the mythical tragelaph which was imagined to be half-goat, half-stag. The root words come from Greek, with τράγος (trágos) meaning "male goat" and έλαφος (élaphos) meaning "stag". The suffix "–ini" refers to their ranking as a tribe.
The alternative name "Strepsicerotini" was published by another British zoologist John Edward Gray as "Strepsiceriae" earlier in 1846. [3] It comes from Greek with στρεπτός (streptós) meaning "twisted" and κέρατος (kératos) meaning "horn", referring to the shape of this group's horns. However, "Strepsiceriae" had the incorrect prefix and suffix, which this was corrected to "Strepsicerotini" in 1945 by American paleontologist George Gaylord Simpson. [14] While the name "Strepsicerotini" was published first, most scientists used the latter name "Tragelaphini" as it is more widely used. [2] [3]
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Phylogenetic relationships of the Bovinae (Bibi et al., 2013) [15] |
The spiral-horned antelopes belong to the subfamily Bovinae which also includes oxen of the tribe Bovini and two aberrant species of Asian antelope, the four-horned antelope and the nilgai which belong to the tribe Boselaphini. The relationship between the tribes varies in research concerning their phylogeny. Most molecular research supports a Bovini and Tragelaphini subclade of Bovinae. [16] [15] [4] There are also some morphological support for oxen being the closet living relatives to the spiral-horned antelopes, most notably both groups have horn cores with a pedicle (the attachment point to the skull). [17]
There has been a few studies which have supported alternate arrangements, [18] [19] [20] one being a sister relationship between the nilgai (Boselaphus tragocamelus) and the spiral-horned antelopes. Historically, the nilgai was classified as a tragelaphine by some scientists. [2] Benirschke et al. (1980) worked on karyotypes with the lesser kudu and found them to share with the nilgai an X chromosome fused with autosome 14. [21] Another study that analyzed the COII gene found the nilgai and lesser kudu being sister species (although the support was low and had limited species sampling). [22] The majority of other molecular and morphological work do not support the placement of the nilgai in Tragelaphini, as it contradicts with their data and results. [16] [17] [15]
Tragelaphini has diverged from their closet sister taxon for the last 15 to 18 million years. [1] It was once thought that spiral-horned antelopes were uniquely African, but there have been Eurasian fossils found in Greece and the Caucasus. [8] Kostopoulos and Koufos (2006) have described Pheraios chryssomallos from fossils found in the Turolian locality of Thessaly, Greece. [8] The authors have found cladistic support of P. chryssomallos being the basal most tragelaphin (or at least a sister taxon to the group) based on 46 cranial features. [8] This suggests that the ancestor of all known species of spiral-horned antelope must have originated in Europe during the late Miocene. [8] The eastern European genus Pontoceros is another example, although this animal has been found in early Pleistocene beds of the Mygdonia basin. [7] This suggests that spiral-horned antelopes have emigrated from Africa and into Eurasia during a latter period in their evolutionary history. [7] In addition, undescribed fossils found in South Asia could be related to spiral-horned antelopes. [1]
Fossils from Africa have been recovered in places such as Olduvai Gorge in Tanzania, but they are sometimes consist of fragmented pieces of leg bone and horn. [17] The oldest known of these species is Tragelaphus moroitu which has been found in the late Miocene to early Pliocene deposits of the Asa Koma, Kuseralee and the Middle Awash deposits in the horn of Africa. [23] [24] Similar in appearance to the nyala (Tragelaphus angasii), T. moroitu was a small antelope and has primitive characteristics in the horn core, making it the most primitive known species of spiral-horn antelope in Africa. [24] The evolution of spiral-horned antelopes based on the abundance of fossils shows they were among the most common species of antelope in Africa, and as climatic changes have occurred throughout their evolution, they have undergone faunal turnovers and adapting to new environments. [24]
Below is the list of fossil species that have been described so far listed in alphabetical order: [8] [23] [24]
Phylogenetic relationships from combined analysis of all molecular data (Willows-Munro et al., 2005) [5] |
Traditionally the tribe was divided into two genera and seven species as shown below: [1]
However recent molecular studies have shown a different arrangement. According to the molecular work on the nuclear DNA and mitochondrial DNA, the lesser kudu and the nyala are the basal most living species. [5] [1] Then around 10 million years ago there was a split between forest-dwelling, gracile species and larger, open-plains species. [5] [1] This renders Tragelaphus paraphyletic in respect to Taurotragus, which either means to make the latter genus into a junior synonym of the former genus or split Tragelaphus into several genera to maintain Taurotragus. [5] In a 2011 large scale taxonomic reorganisation of ungulates by Groves and Grubb, they recognised new cryptic species, and split the traditional species into several geographical species. [6] This was based on morphological comparisons between the species (in particular the dimensions of the skull elements and skin colouration) and defined species based on phylogenetic species concept. [6] Groves and Grubb (2011) furthermore establish of several new genera in light of Tragelaphus being paraphyletic. [6] They have a slight different arrangement of the species in Tragelaphussensu stricto than suggested by genetics. [6] Other scientists have mixed reactions to this proposal, with some preferring the traditional number of species to be seven [25] and others preferring the new taxonomy from Groves and Grubb (2011). [4]
Below is the alternative taxonomy based on Groves and Grubb (2011), [6] with species and subspecies names following Castelló (2016) from Bovids of the World: [4]
The spiral-horned antelopes are medium to large antelopes and generally are tall and long-legged. Horns are present in the males of all species, while females lack them with the exception of bongos and elands. [26] [1] [4] The horns primarily role is used for defensive horn-wrestling which are common during territorial disputes, as well has for horn soiling and sexual display. [26] [1] Other examples of extreme sexual dimorphism can be seen among spiral-horned antelopes. Males are much larger in size than the females, and both sexes have very contrasting pelage coloration. [1] The coloration in females is generally tan to red-brown. Males area a darker hue of the female coloration, which they darken as the animal gets older. [26] Regardless of the coloration of the sexes, most species of spiral-horned antelopes have disruptive coloration that is a distinctive white vertical barring (striped pattern). A spotting pattern can occur as well. [26] [1] Both patterns are individually and geographically variable. These patterns help them to conceal themselves in the dense vegetation from predators. [26] [1] Most species have a white chevron between the eyes, spotted cheeks, the throat having a white patch and the upper-forelegs with dark garters. [26] All except the nyala and the greater kudu have a chest crescent. [26] In addition there are also physical attributes that serve a purpose in social communication such as dorsal crests, white scuts and white tips on the horns. These features help express the animal's emotions as well as alerting members of the herd from predators. [26] [1] Underneath the hair fringe in the hind feet are special glands that encircles the false hooves. [26]
All species are found in a majority of sub-Saharan Africa in various woodland habitats such as rainforests, swamps, open savannahs, mountains, and sub-deserts. [26] With the exception of the eland and sitatunga, spiral-horned antelopes are all browsers. All species forage on green foliage as well as rely on them for cover. [26] During the dry seasons their diet consists mostly foliage from bushes and trees, shoots, twigs and herbs. They also supplement fruit, flowers, and fresh grass when the rainy season arrives. In response to living in dense cover, spiral-horned antelopes are able to move through the thickett by bounding, rushing, and dodging through the vegetation. [26] Most species are water dependent though kudus and elands do not rely on it as they live in more arid environments. [26] Unlike most species of bovids, spiral-horned antelopes are primarily nocturnal though some species can be seen in the early morning and late afternoon. [26] As with all species of antelope, spiral-horned antelopes (especially the young, the old and the weak) are susceptible to some of Africa's major predators, such as lions, leopards, cheetahs, spotted hyenas, and African wild dogs. [26]
When alert to the presence of danger, the necks of these bovines are raised as high as possible. They move in a "goatlike" gait, as they move their neck in a pecking motion (moving the head backwards and forwards). [26] As they spot the predator they immediately stop. The ultimate antipredator strategy is concealment in dense, tall bush, enough for the animal to perfectly camouflage in the vegetation. Sometimes they would raise one leg. [26] If that fails they would sprint away in abrupt flight. They are not known to have high stamina for running, although they are known to jump incredibly high. [26]
Given the various species of spiral-horned antelope the social organization varies between. The extreme ends of the spectrum are the predominately solitary bushbuck and at the other end the highly social giant eland. [26] In general the herd size of spiral-horned antelopes are not large as they rarely exceed more than a few dozen individuals, and are mostly sedentary in nature. [26] [1] The herd composition is unisexual which is mostly females and their young. It is currently believed to be the reason why some spiral-horned antelopes practicing herd as a defense mechanism to protect the young from predators. [26] [1] In the bongo and eland species this led to the development of larger sizes in the females and horns, along with intense social hierarchical ranking (which is not seen in other species which it is only the larger males that dominate over the smaller females). [26] [1] With the exception of the mother and her calf there are no strong social bonds in the herd. [26] [1] [4] These herds are sometimes very open and loosely defined as sometimes females come and go. [26] Males born in the herd will remain in the herd until they developed their primary and secondary sexual characteristics. [26] [4] Once they have developed these traits males leave the herd and become nomads.
It is only during the breeding season males congregate around a female in estrus for a few hours. [26] [1] [4] All spiral-horned antelope species are polyandrous. Aggression is very low among individuals of both sexes though intraspecific competition does occur. In females often they neck-fight as well poke and snap at each other. [26] Males would use their horns for wrestling where they try to attack the face. [26] Sometimes there is even intimation from the males to persuade the female into submission as often she would try and retaliate. [26] The gestation period occurs once the female has been inseminated from the male successfully. In most species it lasts approximately seven months, although in the larger species this extends to eight or nine months. [26] [4] They only give birth to a single calf. Once the young are born, they will not wean until they are around five to six months of age. [4] Females sexually mature by two to three years while for males it is four to five years. [4]
The chromosome number of various species varies. This correlates nicely with the evolutionary relationships among the spiral-horned antelopes. The ancestral Y chromosome was subacrocentric but a pericentric inversion occurred making it submetacentric after the separation of the lesser kudu and nyala. [21] Ancient hybridization also played a key part in the evolution of the chromosomes in all species. [21] Below is a listing of the diploid number 2n as follows (male/female): [21] [27]
Hybridization between bongos and sitatunga produce fertile offspring known as "bongsis" and are well documented. [28] [29] [21] [27] These hybrid animals have a diploid number 2n = 33 and all known cases produced female animals, which had developed horns like female bongos while their stripping pattern was intermediate between bongos and sitatungas with an overall orange coloring. [28] [29] The hybridization of these species adds further support in the bongo and sitatunga being each other's closet relatives. [28] [29] Other hybrids have been documented, such as a male common eland x greater kudu resulting from an accidental crossing from the San Diego Zoo Safari Park in the 1970s. [30] Notable mixed inherited traits were pointed ears as the eland's, but a bit widened like kudu's. The tail was half the length of that of an eland, with a terminal tuft of hair as in kudu. [30] The individual was, however, azoospermic as analysis showed that it completely lacked germ cells, which produce gametes. [30]
Spiral-horned antelopes as a whole are not an endangered group of mammals. [1] [4] Most species are recognized by the IUCN Red List as "least concern", with only the giant eland and mountain nyala being classified as vulnerable [31] and endangered [32] respectively. In addition some local populations and subspecies of the other species have been declining. [1] The main threats to the survival of spiral-horned antelope is habitat destruction and being hunted for bush meat, although most species live in habitat unsuitable for agriculture. [1] In fact in some cases some species might be expanding their numbers (although the accurate population estimates in some species is unknown). [4]
The two eland species are now semi-domesticated as their docile dispositions, large quantities of meat, and resilience in Africa's harsh environments has made them ideal alternatives to cattle in some parts in Africa. [10] [11] [12] [13] In a 2014 comparison between the quality of meat from common elands and cattle, it was found that the meat from male elands was low-fat and more beneficial for human nutritional health but it lacks the sensory texture and flavor that makes cattle meat so appealing to consumers. [13] In addition to being on farm as exotic livestock, elands as well as the other species of spiral-horned antelope have also been brought to private game reserves across the planet (in particular parts in North America and Africa) for recreational hunting. [9] [1] [4] Unlike eland however, most of the other species are not as docile or tolerant of humans as they avoid any contact as possible. [26] [9] [1]
The bongo is a large, mostly nocturnal, forest-dwelling antelope, native to sub-Saharan Africa. Bongos are characterised by a striking reddish-brown coat, black and white markings, white-yellow stripes, and long slightly spiralled horns. It is the only tragelaphid in which both sexes have horns. Bongos have a complex social interaction and are found in African dense forest mosaics. They are the third-largest antelope in the world.
The term antelope refers to numerous extant or recently extinct species of the ruminant artiodactyl family Bovidae that are indigenous to most of Africa, India, the Middle East, Central Asia, and a small area of Eastern Europe. Antelopes do not form a monophyletic group, as some antelopes are more closely related to other bovid groups, like bovines, goats, and sheep, than to other antelopes.
Bovines comprise a diverse group of 10 genera of medium to large-sized ungulates, including cattle, bison, African buffalo, water buffalos, and the four-horned and spiral-horned antelopes. The evolutionary relationship between the members of the group is still debated, and their classification into loose tribes rather than formal subgroups reflects this uncertainty. General characteristics include cloven hooves and usually at least one of the sexes of a species having true horns. The largest extant bovine is the gaur.
The lowland nyala or simply nyala, is a spiral-horned antelope native to southern Africa. It is a species of the family Bovidae and genus Tragelaphus, previously placed in genus Nyala. It was first described in 1849 by George French Angas. The body length is 135–195 cm (53–77 in), and it weighs 55–140 kg (121–309 lb). The coat is maroon or rufous brown in females and juveniles, but grows a dark brown or slate grey, often tinged with blue, in adult males. Females and young males have ten or more white stripes on their sides. Only males have horns, 60–83 cm (24–33 in) long and yellow-tipped. It exhibits the highest sexual dimorphism among the spiral-horned antelopes. It is not to be confused with the endangered mountain nyala living in the Bale region of Ethiopia.
The kudus are two species of antelope of the genus Tragelaphus:
The four-horned antelope, or chousingha, is a small antelope found in India and Nepal. Its four horns distinguish it from most other bovids, which have two horns. The sole member of the genus Tetracerus, the species was first described by French zoologist Henri Marie Ducrotay de Blainville in 1816. Three subspecies are recognised. The four-horned antelope stands nearly 55–64 centimetres (22–25 in) at the shoulder and weighs nearly 17–22 kilograms (37–49 lb). Slender with thin legs and a short tail, the four-horned antelope has a yellowish brown to reddish coat. One pair of horns is located between the ears, and the other on the forehead. The posterior horns are always longer than the anterior horns, which might be mere fur-covered studs. While the posterior horns measure 8–12 centimetres (3.1–4.7 in), the anterior ones are 2–5 centimetres (0.79–1.97 in) long.
The saola, also called spindlehorn, Asian unicorn, or infrequently, Vu Quang bovid, is one of the world's rarest large mammals, a forest-dwelling bovine native to the Annamite Range in Vietnam and Laos. It was described in 1993 following a discovery of remains in Vũ Quang National Park by a joint survey of the Vietnamese Ministry of Forestry and the World Wide Fund for Nature. Saolas have since been kept in captivity multiple times, although only for short periods as they died within a matter of weeks to months. The species was first reported in 1992 by Do Tuoc, a forest ecologist, and his associates. The first photograph of a living saola was taken in captivity in 1993. The most recent one was taken in 2013 by a movement-triggered camera in the forest of central Vietnam. It is the only species in the genus Pseudoryx.
Tragelaphus is a genus of medium-to-large-sized spiral-horned antelopes. It contains several species of bovines, all of which are relatively antelope-like. Species in this genus tend to be large in size and lightly built, and have long necks and considerable sexual dimorphism. Elands, including the common eland, are embedded within this genus, meaning that Taurotragus must be subsumed into Tragelaphus to avoid paraphyly. Alternatively, Taurotragus could be maintained as a separate genus, if the nyala and the lesser kudu are relocated to their own monospecific genera, respectively Nyala and Ammelaphus. Other generic synonyms include Strepsiceros and Boocercus. The name "Tragelaphus" comes from the mythical tragelaph.
The sitatunga or marshbuck is a swamp-dwelling medium-sized antelope found throughout central Africa, centering on the Democratic Republic of the Congo, the Republic of the Congo, Cameroon, parts of Southern Sudan, Equatorial Guinea, Burundi, Ghana, Botswana, Rwanda, Zambia, Gabon, the Central African Republic, Tanzania, Uganda and Kenya. The sitatunga is mostly confined to swampy and marshy habitats. Here they occur in tall and dense vegetation as well as seasonal swamps, marshy clearings in forests, riparian thickets and mangrove swamps.
The Cape bushbuck, also known as imbabala is a common, medium-sized bushland-dwelling, and a widespread species of antelope in sub-Saharan Africa. It is found in a wide range of habitats, such as rain forests, montane forests, forest-savanna mosaic, savanna, bushveld, and woodland. Its stands around 90 cm (35 in) at the shoulder and weigh from 45 to 80 kg. They are generally solitary, territorial browsers.
The mountain nyala or balbok, is a large antelope found in high altitude woodlands in a small part of central Ethiopia. It is a monotypic species first described by English naturalist Richard Lydekker in 1910. The males are typically 120–135 cm (47–53 in) tall while females stand 90–100 cm (35–39 in) at the shoulder. Males weigh 180–300 kg (400–660 lb) and females weigh 150–200 kg (330–440 lb). The coat is grey to brown, marked with two to five poorly defined white strips extending from the back to the underside, and a row of six to ten white spots. White markings are present on the face, throat and legs as well. Males have a short dark erect crest, about 10 cm (3.9 in) high, running along the middle of the back. Only males possess horns.
The lesser kudu is a medium-sized bushland antelope found in East Africa. The species is a part of the ungulate genus Tragelaphus, along with several other related species of striped, spiral-horned African bovids, including the related greater kudu, the bongo, bushbuck, common and giant elands, nyala and sitatunga. It was first scientifically described by English zoologist Edward Blyth (1869).
The greater kudu is a large woodland antelope, found throughout eastern and southern Africa. Despite occupying such widespread territory, they are sparsely populated in most areas due to declining habitat, deforestation, and poaching. The greater kudu is one of two species commonly known as kudu, the other being the lesser kudu, T. imberbis.
The common eland, also known as the southern eland or eland antelope, is a large-sized savannah and plains antelope found in East and Southern Africa. It is a species of the family Bovidae and genus Taurotragus. An adult male is around 1.6 m (5.2 ft) tall at the shoulder and can weigh up to 942 kg (2,077 lb) with a typical range of 500–600 kg (1,100–1,300 lb), 340–445 kg (750–981 lb) for females). It is the second-largest antelope in the world, being slightly smaller on average than the giant eland. It was scientifically described by Peter Simon Pallas in 1766.
The giant eland, also known as the Lord Derby's eland and greater eland, is an open-forest and savanna antelope. A species of the family Bovidae and genus Taurotragus, it was described in 1847 by John Edward Gray. The giant eland is the largest species of antelope, with a body length ranging from 220–290 cm (87–114 in). There are two subspecies: T. d. derbianus and T. d. gigas.
Taurotragus is a genus of large antelopes of the African savanna, commonly known as elands. It contains two species: the common eland T. oryx and the giant eland T. derbianus.
The tribe Bovini, or wild cattle, are medium to massive bovines that are native to Eurasia, North America, and Africa. These include the enigmatic, antelope-like saola, the African and Asiatic buffalos, and a clade that consists of bison and the wild cattle of the genus Bos. Not only are they the largest members of the subfamily Bovinae, they are the largest species of their family Bovidae. The largest species is the gaur, weighing up to 1,500 kg (3,300 lb).
Elaeophora sagitta is a parasitic nematode found in the heart, coronary arteries and pulmonary arteries of several ruminant species and African buffaloes in Africa. Infestation usually occurs without significant health effects in the Greater kudu, but may affect cardiac function in some other host species.
The northern bushbuck or harnessed bushbuck, is a medium-sized antelope, widespread in sub-Saharan-Africa. The northern bushbuck species has been separated from the Cape bushbuck, a southern and eastern species.
The Albany thickets is an ecoregion of dense woodland in southern South Africa, which is concentrated around the Albany region of the Eastern Cape.