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The hartebeest (Template:IPAc-en;<ref name=MW/> Alcelaphus buselaphus), also known as kongoni or kaama, is an African antelope. It is the only member of the genus Alcelaphus. Eight subspecies have been described, including two sometimes considered to be independent species. A large antelope, the hartebeest stands just over Template:Convert at the shoulder, and has a typical head-and-body length of Template:Convert. The weight ranges from Template:Convert. It has a particularly elongated forehead and oddly-shaped horns, a short neck, and pointed ears. Its legs, which often have black markings, are unusually long. The coat is generally short and shiny. Coat colour varies by the subspecies, from the sandy brown of the western hartebeest to the chocolate brown of the Swayne's hartebeest. Both sexes of all subspecies have horns, with those of females being more slender. Horns can reach lengths of Template:Convert. Apart from its long face, the large chest and the sharply sloping back differentiate the hartebeest from other antelopes. A conspicuous hump over the shoulders is due to the long dorsal processes of the vertebrae in this region.<ref>Template:Cite book</ref>
Gregarious animals, hartebeest form herds of 20 to 300 individuals. They are very alert and non-aggressive. They are primarily grazers, with their diets consisting mainly of grasses. Mating in hartebeest takes place throughout the year with one or two peaks, and depends upon the subspecies and local factors. Both males and females reach sexual maturity at one to two years of age. Gestation is eight to nine months long, after which a single calf is born. Births usually peak in the dry season. The lifespan is 12 to 15 years.
Inhabiting dry savannas and wooded grasslands, hartebeest often move to more arid places after rainfall. They have been reported from altitudes on Mount Kenya up to Template:Convert. The hartebeest was formerly widespread in Africa, but populations have undergone a drastic decline due to habitat destruction, hunting, human settlement, and competition with livestock for food. Each of the eight subspecies of the hartebeest has a different conservation status. The Bubal hartebeest was declared extinct by the International Union for Conservation of Nature (IUCN) in 1994. While the populations of the red hartebeest are on the rise, those of the Tora hartebeest, already Critically Endangered, are falling. The hartebeest is extinct in Algeria, Egypt, Lesotho, Libya, Morocco, Somalia, and Tunisia; but has been introduced into Eswatini and Zimbabwe. It is a popular game animal due to its highly regarded meat.
EtymologyEdit
The vernacular name "hartebeest" may have originated from the obsolete Dutch word hertebeest,<ref name=mares>Template:Cite book</ref> literally deer beast,<ref name=MW>Template:MerriamWebsterDictionary</ref> based on the resemblance (to early Dutch settlers) of the antelope to deer.<ref name=Llewellyn1936>Template:Cite book</ref> The first use of the word "hartebeest" in South African literature was in Dutch colonial administrator Jan van Riebeeck's journal Daghregister in 1660. He wrote: "Meester Pieter ein hart-beest geschooten hadde (Master Pieter [van Meerhoff] had shot one hartebeest)".<ref name=skinner>Template:Cite book</ref> Another name for the hartebeest is kongoni,<ref name=kingdon2013/> a Swahili word.<ref name=MW2>Template:MerriamWebsterDictionary</ref> Kongoni is often used to refer in particular to one of its subspecies, Coke's hartebeest.<ref name="swank">Template:Cite book</ref>
TaxonomyEdit
The scientific name of the hartebeest is Alcelaphus buselaphus. First described by German zoologist Peter Simon Pallas in 1766, it is classified in the genus Alcelaphus and placed in the family Bovidae.<ref name=MSW3>Template:Cite book</ref> In 1979, palaeontologist Elisabeth Vrba supported Sigmoceros as a separate genus for Lichtenstein's hartebeest, a kind of hartebeest, as she assumed it was related to Connochaetes (wildebeest).<ref name=nowak>Template:Cite book</ref><ref name=flagstad2001/> She had analysed the skull characters of living and extinct species of antelope to make a cladogram, and argued that a wide skull linked Lichtenstein's hartebeest with Connochaetes.<ref>Template:Cite journal</ref> However, this finding was not replicated by Alan W. Gentry of the Natural History Museum, who classified it as an independent species of Alcelaphus.<ref>Template:Cite book</ref> Zoologists such as Jonathan Kingdon and Theodor Haltenorth considered it to be a subspecies of A. buselaphus.<ref name=MSW3/> Vrba dissolved the new genus in 1997 after reconsideration.<ref name=groves>Template:Cite book</ref> An MtDNA analysis could find no evidence to support a separate genus for Lichtenstein's hartebeest. It also showed the tribe Alcelaphini to be monophyletic, and discovered close affinity between the Alcelaphus and the sassabies (genus Damaliscus)—both genetically and morphologically.<ref name=matthee>Template:Cite journal</ref>
SubspeciesEdit
Eight subspecies are identified, of which two – A. b. caama and A. b. lichtensteinii – have been considered to be independent species. However, a 1999 genetic study by P. Arctander of the University of Copenhagen and colleagues, which sampled the control region of the mitochondrial DNA, found that these two formed a clade within A. buselaphus, and that recognising these as species would render A. buselaphus paraphyletic (an unnatural grouping). The same study found A. b. major to be the most divergent, having branched off before the lineage split to give a combined caama/lichtensteinii lineage and another that gave rise to the remaining extant subspecies.<ref name=arctander>Template:Cite journal</ref> Conversely a 2001 phylogenetic study, based on D–loop and cytochrome b analysis by Øystein Flagstad (of the Norwegian Institute for Nature Research, Trondheim) and colleagues, found that the southern lineage of A. b. caama and A. lichtensteinii diverged earliest.<ref name=flagstad2001/> Analysis of skull structure supports partition into three major divisions: A. b. buselaphus division (nominate, also including A. b. major division), A. b. tora division (also including A. b. cokii and A. b. swaynei) and A. b. lelwel division.<ref name=MSW3/> Another analysis of cytochrome b and D-loop sequence data shows a notable affinity between the A. b. lelwel and A. b. tora divisions.<ref name=flagstad2000>Template:Cite journal</ref>
The eight subspecies, including the two controversial ones, are:<ref name=iucn/><ref name=ITIS>{{#if:625077 | {{#invoke:template wrapper|wrap|_template=cite web|_exclude=id,ID,taxon
| url = https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=625077 | title = Alcelaphus buselaphus | publisher = Integrated Taxonomic Information System }}
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- † A. b. buselaphus (Pallas, 1766) : Known as the bubal hartebeest or northern hartebeest. Formerly occurred across northern Africa, from Morocco to Egypt. It was exterminated by the 1920s.<ref name=east/> It was declared extinct in 1994 by the International Union for the Conservation of Nature and Natural Resources (IUCN).<ref name=bubal>Template:Cite iucn</ref><ref name=mallon>Template:Cite book</ref>
- A. b. caama (Saint-Hilaire, 1803) : Known as the red hartebeest or Cape hartebeest. Formerly occurred in southern Angola; northern and eastern savannahs of Namibia; central, southern and southwestern Botswana; Northern Cape, Eastern Cape, Western Cape, Free State, Northwest and Gauteng provinces and western KwaZulu-Natal of South Africa. Presently has been eliminated from all these areas except Northern Cape, central and southwestern Botswana and Namibia. Major re-introductions have taken place in these countries.<ref name=east/> The population of this hartebeest is on the rise.<ref name=caama>Template:Cite iucn</ref>
- A. b. cokii Günther, 1884: Known as Coke's hartebeest or kongoni. Native to and confined within Kenya and northern Tanzania.<ref name=east/>
- A. b. lelwel (Heuglin, 1877) : Known as the Lelwel hartebeest. Formerly found in northern and northeastern Democratic Republic of the Congo; southeastern and southwestern Sudan; and the northwestern extreme of Tanzania.<ref name=east>Template:Cite book</ref> Drastic population decrease since the 1980s has confined most individuals to protected areas inside and outside its range.<ref name=lelwel>Template:Cite iucn</ref>
- A. b. lichtensteinii (Peters, 1849) : Known as Lichtenstein's hartebeest. Inhabits the miombo woodlands of eastern and southern Africa.<ref name=rafferty1>Template:Cite book</ref> It is native to Angola, the Democratic Republic of Congo, Malawi, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe.<ref name="lh"/>
- A. b. major (Blyth, 1869) : Known as the western hartebeest. Formerly occurred widely in Mali, Niger, Senegal, Gambia, Guinea-Bissau, Guinea, Ivory Coast, Ghana, Nigeria, southwestern Chad, Cameroon, western Central African Republic and Benin. Nowadays it occurs in much lower numbers mainly in protected areas of these countries. It is probably extinct in Gambia.<ref name=east/>
- A. b. swaynei (Sclater, 1892) : Known as Swayne's hartebeest. Restricted to the southern Rift Valley in Ethiopia, confirmed and listed as an endemic species. It formerly occurred throughout the Rift Valley, and its range extended eastward into northwestern Somalia. It disappeared from Somalia by 1930 due to the outcome of the rinderpest epidemic, illegal poaching, and livestock overgrazing.<ref name=east/> Confirmed as endangered species, its populations are low with at least 1500 individuals, and on verge of population decline.<ref name=lewis>Template:Cite journal</ref><ref>Tamrat, M., Atickem, A., Flagstad, Ø, Fischer, M., Roos, C., Evangelista, P., . . . Zinner, D. (2022). Swayne's hartebeest in Ethiopia: Population estimate, genetic variability and competition with livestock. Oryx, 56(3), 336-344. doi:10.1017/S0030605320000927</ref>
- A. b. tora Gray, 1873 : Known as the tora hartebeest. Formerly occurred in northwestern Ethiopia and western and southwestern Eritrea.<ref name=hildyard/> Its present status is unclear, though locals have reported small numbers from these areas.<ref name=east
/>
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Genetics and hybridsEdit
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In 2000, a study scrutinised two major populations of the Swayne's hartebeest, from the Senkele Wildlife Sanctuary and the Nechisar National Park, for mitochondrial (D-loop) and nuclear (microsatellite) variability in an attempt to estimate the levels of genetic variation between the populations and within the subspecies. The results showed a remarkable differentiation between the two populations; that from the Senkele Wildlife Sanctuary showed more genetic diversity than the one from the Nechisar National Park. Another revelation was that the translocation of the individuals from the Senkele Wildlife Sanctuary in 1974 had not made a significant contribution to the gene pool of the Nechisar National Park. Additionally, the Swayne hartebeest populations were compared with a large red hartebeest population, and both subspecies were found to have a high degree of genetic variation. The study advocated in situ conservation of the Swayne's hartebeest and a renewed attempt at its translocation in order to conserve genetic diversity and increase its population in both the protected areas.<ref name=flagstad2000/>
The diploid number of chromosomes in the hartebeest is 40. Hybrids are usually reported from areas where ranges of two subspecies overlap.<ref name=kingdon2013/> Hybrids between the Lelwel and Tora hartebeest have been reported in eastern Sudan and western Ethiopia, in a stretch southward from the Blue Nile to about 9° N latitude.<ref name=lelwelsci/> A study proved a male hybrid of the red hartebeest and the blesbok (Damaliscus pygargus) to be sterile. Sterility of the hybrid was attributed to difficulties in segregation during meiosis, indicated by azoospermia and a low number of germ cells in its seminiferous tubules.<ref name=robinson>Template:Cite journal</ref>
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There are three well-defined hybrids between the subspecies:
- Alcelaphus lelwel x cokii: Known as the Kenya Highland hartebeest or the Laikipia hartebeest. It is a cross between the Lelwel and Coke's hartebeest.<ref name=castello/> This hybrid is lighter in colour and larger than Coke's hartebeest. It is a light buff with reddish-tawny upper parts, and the head is longer than in Coke's hartebeest. Both sexes have horns, which are heavier as well as longer than those of the parents. It was formerly distributed throughout the western Kenyan highlands, between Lake Victoria and Mount Kenya, but is now believed to be restricted to the Lambwe Valley (south-west Kenya) and Laikipia and nearby regions of west-central Kenya.<ref name=Augustine2011>Template:Cite book</ref><ref name=kenyasci>{{#invoke:citation/CS1|citation
|CitationClass=web }}</ref>
- The Jackson's hartebeest does not have a clear taxonomic status. Like the form above, it is regarded as a hybrid between the Lelwel and Coke's hartebeest, and has a similar distribution. The African Antelope Database (1998) treats it as synonymous to the Lelwel hartebeest.<ref name=east/> From Lake Baringo to Mount Kenya, the Jackson's hartebeest significantly resembles the Lelwel hartebeest, whereas from Lake Victoria to the southern part of the Rift Valley it tends to be more like the Coke's hartebeest.<ref name=ruxton/>
- Alcelaphus lelwel x swaynei : Also known as the Neumann's hartebeest, named after traveller and hunter Arthur Henry Neumann.<ref name=ruxton>Template:Cite journal</ref> This is considered to be a cross between the Lelwel hartebeest and Swayne's hartebeest.<ref name=castello>Template:Cite book</ref> The face is longer than that of the Swayne's hartebeest. The colour of the coat is a golden brown, paler towards the underparts. The chin has a hint of black and the tail ends in a black tuft. Both sexes have longer horns than the Swayne's hartebeest. The horns grow in a wide "V" shape, unlike the wide bracket shape of Swayne's hartebeest and the narrow "V" of Lelwel hartebeest, curving backward and slightly inward. It occurs in Ethiopia, in a small area to the east of Omo River and north of Lake Turkana, stretching north-east of Lake Chew Bahir to near Lake Chamo.<ref name=neumannsci>{{#invoke:citation/CS1|citation
|CitationClass=web }}</ref>
EvolutionEdit
The genus Alcelaphus emerged about 4.4 million years ago in a clade whose other members were Damalops, Numidocapra, Rabaticeras, Megalotragus, Oreonagor, and Connochaetes. An analysis using phylogeographic patterns within hartebeest populations suggested a possible origin of Alcelaphus in eastern Africa.<ref name=Harris>Template:Cite book</ref> Alcelaphus quickly radiated across the African savannas, replacing several previous forms (such as a relative of the hirola). Flagstad and colleagues showed an early split in the hartebeest populations into two distinct lineages around 0.5 million years ago – one to the north and the other to the south of the equator. The northern lineage further diverged into eastern and western lineages, nearly 0.4 million years ago, most probably as a result of the expanding central African rainforest belt and subsequent contraction of savanna habitats during a period of global warming. The eastern lineage gave rise to the Coke's, Swayne's, Tora and Lelwel hartebeest; and from the western lineage evolved the Bubal and western hartebeest. The southern lineage gave rise to Lichtenstein's and red hartebeest. These two taxa are phylogenetically close, having diverged only 0.2 million years ago. The study concluded that these major events throughout the hartebeest's evolution are strongly related to climatic factors, and that there had been successive bursts of radiation from a more permanent population—a refugium—in eastern Africa; this could be vital to understanding the evolutionary history of not only the hartebeest but also other mammals of the African savanna.<ref name=flagstad2001>Template:Cite journal</ref>
The earliest fossil record dates back to nearly 0.7 million years ago.<ref name=kingdon2013>Template:Cite book</ref> Fossils of the red hartebeest have been found in Elandsfontein, Cornelia (Free State) and Florisbad in South Africa, as well as in Kabwe in Zambia.<ref name=berger>Template:Cite book</ref> In Israel, hartebeest remains have been found in northern Negev, Shephelah, Sharon Plain and Tel Lachish. This population of the hartebeest was originally limited to the open country of the southernmost regions of the southern Levant. It was probably hunted in Egypt, which affected the numbers in the Levant, and disconnected it from its main population in Africa.<ref name=tsahar>Template:Cite journal Template:Open access</ref>
DescriptionEdit
A large antelope with a particularly elongated forehead and oddly shaped horns, the hartebeest stands just over Template:Convert at the shoulder, and has a typical head-and-body length of Template:Convert. The weight ranges from Template:Convert. The tail, Template:Convert long, ends in a black tuft.<ref name=kingdon>Template:Cite book</ref> The other distinctive features of the hartebeest are its long legs (often with black markings), short neck, and pointed ears.<ref name=Macdonald>Template:Cite book</ref> A study correlated the size of hartebeest species to habitat productivity and rainfall.<ref name=capellini>Template:Cite journal</ref> The western hartebeest is the largest subspecies, and has a characteristic white line between the eyes.<ref name=westernsci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The red hartebeest is also large, with a black forehead and a contrasting light band between the eyes.<ref name=redsci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The large Lelwel hartebeest has dark stripes on the front of its legs.<ref name=lelwelsci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Coke's hartebeest is moderately large, with a shorter forehead and longer tail in comparison to the other subspecies.<ref name=cokesci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Lichtenstein's hartebeest is smaller, with dark stripes on the front of the legs, as in the Lelwel hartebeest.<ref name=Lsci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The Swayne's hartebeest is smaller than the Tora hartebeest, but both have a shorter forehead and similar appearance.<ref name=swaynesci>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
Generally short and shiny, the coat varies in colour according to subspecies.<ref name=estes/> The western hartebeest is a pale sandy-brown, but the front of the legs are darker.<ref name=westernsci/> The red hartebeest is a reddish-brown, with a dark face. Black markings can be observed on the chin, the back of the neck, shoulders, hips and legs; these are in sharp contrast with the broad white patches that mark its flanks and lower rump.<ref name=redsci/><ref name=wwsa>Template:Cite book</ref> The Lelwel hartebeest is a reddish tan.<ref name=lelwelsci/> Coke's hartebeest is reddish to tawny in the upper parts, but has relatively lighter legs and rump.<ref name=cokesci/> Lichtenstein's hartebeest is reddish brown, though the flanks are a lighter tan and the rump whitish.<ref name=Lsci/> The Tora hartebeest is a dark reddish brown in the upper part of the body, the face, the forelegs and the rump, but the hindlegs and the underbelly are a yellowish white.<ref name=hildyard>Template:Cite book</ref><ref name=heckel>Template:Cite report</ref> The Swayne's hartebeest is a rich chocolate brown with fine spots of white that are actually the white tips of its hairs. Its face is black save for the chocolate band below the eyes. The shoulders and upper part of the legs are black.<ref name=swaynesci/> Fine textured, the body hair of the hartebeest is about Template:Convert long.<ref name="nowak"/> The hartebeest has preorbital glands (glands near the eyes) with a central duct, that secrete a dark sticky fluid in Coke's and Lichtenstein's hartebeest, and a colourless fluid in the Lelwel hartebeest.<ref name=estes/>
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Both sexes of all subspecies have horns, with those of females being more slender. Horns can reach lengths of Template:Convert; the maximum horn length is Template:Convert, recorded from a Namibian red hartebeest.<ref name=kingdon/> The horns of the western hartebeest are thick and appear U-shaped from the front and Z-shaped from the sides, growing backward at first and then forward, ending with a sharp backward turn.<ref name=westernsci/> The horns of the red and the Lelwel hartebeest are similar to those of the western hartebeest, but appear V-shaped when viewed from the front.<ref name=lelwelsci/><ref name=redsci/> The Lichtenstein's hartebeest has thick parallel ringed horns, with a flat base. Its horns are shorter than those of other subspecies, curving upward then sharply forward, followed by an inward turn at an angle of about 45° and a final backward turn.<ref name=Lsci/> The horns of Swayne's hartebeest are thin and shaped like parentheses, curving upward and then backward.<ref name=swaynesci/> The horns of the Tora hartebeest are particularly thin and spread out sideways, diverging more than in any other subspecies.<ref name=heckel/>
Apart from its long face, the large chest and the sharply sloping back differentiate the hartebeest from other antelopes.<ref name=mares/> The hartebeest shares several physical traits with the sassabies (genus Damaliscus), such as an elongated and narrow face, the shape of the horns, the pelage texture and colour, and the terminal tuft of the tail. The wildebeest have more specialised skull and horn features than the hartebeest.<ref name=estes/> The hartebeest exhibits sexual dimorphism, but only slightly, as both sexes bear horns and have similar body masses. The degree of sexual dimorphism varies by subspecies. Males are 8% heavier than females in Swayne's and Lichtenstein's hartebeest, and 23% heavier in the red hartebeest. In one study, the highest dimorphism was found in skull weight.<ref name=capellini2>Template:Cite book</ref> Another study concluded that the length of the breeding season is a good predictor of dimorphism in pedicle (the bony structures from which the horns grow) height and skull weight, and the best predictor of the horn circumference.<ref name=capellini3>Template:Cite journal</ref>
Ecology and behaviourEdit
Active mainly during daytime, the hartebeest grazes in the early morning and late afternoon, and rests in shade around noon. Gregarious, the species forms herds of up to 300 individuals. Larger numbers gather in places with abundant grass. In 1963, a congregation of 10,000 animals was recorded on the plains near Sekoma Pan in Botswana.<ref name=estes/> However, moving herds are not so cohesive, and tend to disperse frequently. The members of a herd can be divided into four groups: territorial adult males, non-territorial adult males, young males, and the females with their young. The females form groups of five to 12 animals, with four generations of young in the group. Females fight for dominance over the herd.<ref name=kingdon/> Sparring between males and females is common.<ref name=kingdon2013/> At three or four years of age, the males can attempt to take over a territory and its female members. A resident male defends his territory and will fight if provoked.<ref name=capellini2/> The male marks the border of his territory through defecation.<ref name=kingdon/>
Hartebeest are remarkably alert and cautious animals with highly developed brains.<ref>Template:Cite journal</ref><ref name=schaller>Template:Cite book</ref> Generally calm in nature, hartebeest can be ferocious when provoked. While feeding, one individual stays on the lookout for danger, often standing on a termite mound to see farther. At times of danger, the whole herd flees in a single file after an individual suddenly starts off.<ref name=kingdon/> Adult hartebeest are preyed upon by lions, leopards, hyenas and wild dogs; cheetahs and jackals target juveniles.<ref name=kingdon/> Crocodiles may also prey on hartebeest.<ref>Template:Cite book</ref>
The thin long legs of the hartebeest provide for a quick escape in an open habitat; if attacked, the formidable horns are used to ward off the predator. The elevated position of the eyes enables the hartebeest to inspect its surroundings continuously even as it is grazing. The muzzle has evolved so as to derive maximum nutrition from even a frugal diet.<ref name=kingdon2013/> The horns are also used during fights among males for dominance in the breeding season;<ref name=capellini3/> the clash of the horns is loud enough that it can be heard from hundreds of metres away.<ref name=kingdon2013/> The beginning of a fight is marked with a series of head movements and stances, as well as depositing droppings on dung piles. The opponents drop onto their knees and, after giving a hammer-like blow, begin wrestling, their horns interlocking. One attempts to fling the head of the other to one side to stab the neck and shoulders with his horns.<ref name=capellini2/> Fights are rarely serious, but can be fatal if they are.<ref name=estes/>
Like the sassabies, hartebeest produce quiet quacking and grunting sounds. Juveniles tend to be more vocal than adults, and produce a quacking call when alarmed or pursued.<ref name=kingdon/> The hartebeest uses defecation as an olfactory and visual display.<ref name=estes/> Herds are generally sedentary, and tend to migrate only under adverse conditions such as natural calamities.<ref name=verlinden>Template:Cite journal</ref> The hartebeest is the least migratory in the tribe Alcelaphini (which also includes wildebeest and sassabies), and also consumes the least amount of water and has the lowest metabolic rate among the members of the tribe.<ref name=estes/>
Parasites and diseasesEdit
Several parasites have been isolated from the hartebeest.<ref name=boomker>Template:Cite journal</ref><ref name=howard/> These parasites regularly alternate between hartebeest and gazelles or wildebeest.<ref name=pester>Template:Cite journal</ref> Hartebeest can be infected with theileriosis due to Rhipicephalus evertsi and Theileria species.<ref>Template:Cite journal</ref> South of the Sahara, common parasites include Loewioestrus variolosus, Gedoelstia cristata and G. hassleri. The latter two species can cause serious diseases such as encephalitis.<ref name=spinage>Template:Cite book</ref> However, parasites are not always harmful – 252 larvae were found in the head of one Zambian individual without any pathogenicity.<ref name=howard>Template:Cite journal</ref> Nematodes, cestodes, paramphistomes; and the roundworm Setaria labiatopapillosa have also been isolated from the hartebeest.<ref name=belem>Template:Cite journal Template:Open access</ref><ref>Template:Cite journal</ref> In 1931, a red hartebeest in Gobabis (southwestern Africa) was infected with long, thin worms. These were named Longistrongylus meyeri after their collector, T. Meyer.<ref>Template:Cite journal</ref>
DietEdit
Hartebeest are primarily grazers, and their diets consist mostly of grasses.<ref name=awf>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> A study in the Nazinga Game Ranch in Burkina Faso found that the hartebeest's skull structure eased the acquisition and chewing of highly fibrous foods.<ref name=schuette/> The hartebeest has much lower food intake than the other members of Alcelaphini. The long thin muzzle of the hartebeest assists in feeding on leaf blades of short grasses and nibbling off leaf sheaths from grass stems. In addition to this, it can derive nutritious food even from tall senile grasses. These adaptations of the hartebeest enable the animal to feed well even in the dry season, which is usually a difficult period for grazers.<ref name=kingdon2013/> For instance, in comparison with the roan antelope, the hartebeest is better at procuring and chewing the scarce regrowth of perennial grasses at times when forage is least available.<ref name=schuette/> These unique abilities could have allowed the hartebeest to prevail over other animals millions of years ago, leading to its successful radiation across Africa.<ref name=kingdon2013/>
Grasses generally comprise at least 80 per cent of the hartebeest's diet, but they account for over 95 per cent of their food in the wet season, October to May. Jasminum kerstingii is part of the hartebeest's diet at the start of the rainy season. Between seasons, they mainly feed on the culms of grasses.<ref name=schuette>Template:Cite journal</ref> A study found that the hartebeest is able to digest a higher proportion of food than the topi and the wildebeest.<ref name=murray>Template:Cite journal</ref> In areas with scarce water, it can survive on melons, roots, and tubers.<ref name=estes/>
In a study of grass selectivity among the wildebeest, zebra, and the Coke's hartebeest, the hartebeest showed the highest selectivity. All animals preferred Themeda triandra over Pennisetum mezianum and Digitaria macroblephara. More grass species were eaten in the dry season than in the wet season.<ref name=casebeer>Template:Cite journal</ref>
ReproductionEdit
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Mating in hartebeest takes place throughout the year, with one or two peaks that can be influenced by the availability of food.<ref name=awf/> Both males and females reach sexual maturity at one to two years of age. Reproduction varies by the subspecies and local factors.<ref name=nowak/> Mating takes place in the territories defended by a single male, mostly in open areas.<ref name=awf/> The males may fight fiercely for dominance,<ref name=capellini2/> following which the dominant male smells the female's genitalia, and follows her if she is in oestrus. Sometimes a female in oestrus holds out her tail slightly to signal her receptivity,<ref name=estes/> and the male tries to block the female's way. She may eventually stand still and allow the male to mount her. Copulation is brief and is often repeated, sometimes twice or more in a minute.<ref name=estes/> Any intruder at this time is chased away.<ref name=kingdon/> In large herds, females often mate with several males.<ref name=estes/>
Gestation is eight to nine months long, after which a single calf weighing about Template:Convert is born. Births usually peak in the dry season, and take place in thickets – unlike the wildebeest, which give birth in groups on the plains.<ref name=estes/> Though calves can move about on their own shortly after birth, they usually lie in the open in close proximity of their mothers.<ref name=castello/> The calf is weaned at four months,<ref name=castello/> but young males stay with their mothers for two and a half years, longer than in other Alcelaphini.<ref name=estes>Template:Cite book</ref> Often the mortality rate of male juveniles is high, as they have to face the aggression of territorial adult males and are also deprived of good forage by them.<ref name=kingdon/> The lifespan is 12 to 15 years.<ref name=awf/>
HabitatEdit
Hartebeest inhabit dry savannas, open plains and wooded grasslands,<ref name=nowak/> often moving into more arid places after rainfall. They are more tolerant of wooded areas than other Alcelaphini, and are often found on the edge of woodlands.<ref name=awf/> They have been reported from altitudes on Mount Kenya up to Template:Convert.<ref name=iucn/> The red hartebeest is known to move across large areas, and females roam home ranges of over Template:Convert, with male territories Template:Convert in size.<ref name=mills>Template:Cite book</ref> Females in the Nairobi National Park (Kenya) have individual home ranges stretching over Template:Convert, which are not particularly associated with any one female group. Average female home ranges are large enough to include 20 to 30 male territories.<ref name=Macdonald/>
Status and conservationEdit
Each hartebeest subspecies is listed under a different conservation status by the International Union for Conservation of Nature. The species as a whole is classified as Least Concern by the IUCN.<ref name=iucn/> The hartebeest is extinct in Algeria, Egypt, Lesotho, Libya, Morocco, Somalia, and Tunisia.<ref name=iucn/>
- The Bubal hartebeest has been declared extinct since 1994.<ref name=bubal/> German explorer Heinrich Barth, in his works of 1857, cites firearms and European intrusion among the reasons for the decrease in its numbers.<ref name=yadav>Template:Cite book</ref> It was extinct in Tunisia by the late 19th century.<ref>Template:Cite book</ref> The last individual was shot in Missour (Algeria) in 1925.<ref name=harper>Template:Cite book</ref>
- Coke's hartebeest is listed as Least Concern. This species has been greatly affected by habitat destruction, and about 42,000 Coke's hartebeest occur today in Mara, Serengeti National Park, and Tarangire National Park in Tanzania and Tsavo East National Park in Kenya. The population is decreasing, and 70% of the population lives in protected areas.<ref name=coke>Template:Cite iucn</ref>
- The Lelwel hartebeest is listed as Endangered, and numbers have declined greatly since the 1980s, when its population was over 285,000. It was formerly distributed mainly in the Central African Republic, Ethiopia, northern and northeastern Democratic Republic of Congo and southern Sudan.<ref name=east/> Fewer than 70,000 individuals are left.<ref name=lelwel/> Most of the population nowadays is found in Chad in the Salamat region and the Zakouma National Park (Chad), the National Park population benefiting from improved protection and seeing an increase in population since the 1980s; Manovo-Gounda St. Floris National Park and Bamingui-Bangoran National Park and Biosphere Reserve in the Central African Republic, where the populations have been falling; Rumanyika Orugundu Game Reserve and Ibanda Game Reserve in Tanzania; and Murchison Falls National Park in Uganda.<ref name=east/>
- Lichtenstein's hartebeest is listed as Least Concern, and occurs in protected areas such as the Selous Game Reserve and in the wild in southern and western Tanzania and Zambia.<ref name=lh>Template:Cite iucn</ref>
- The red hartebeest is listed as Least Concern. It is the most widespread, with increasing numbers after its reintroduction into protected and private areas. However, it has been extinct in Lesotho since the twentieth century.<ref name=east/> Its population is estimated to be over 130,000 (as of 2008),<ref name=caama/> mostly in southern Africa.<ref name=mills/> In Namibia, the largest population occurs in the Etosha National Park. A reintroduced population is flourishing in the Malolotja Nature Reserve (Eswatini), outside its range. However, numbers have seen a sharp fall in southwestern Botswana.<ref name=east/>
- The Tora hartebeest is listed as Critically Endangered; the IUCN has ascertained that fewer than 250 mature individuals survive as of 2008. It is possibly extinct due to excessive hunting and agricultural expansion, but may still exist in smaller numbers in Eritrea and Ethiopia.<ref name=tora>Template:Cite iucn</ref> There have been unconfirmed reports of sightings by locals of the Tora hartebeest southeast of the Dinder National Park, from where it had disappeared before 1960.<ref name=east/>
- Swayne's hartebeest is listed as Endangered, and is close to being Critically Endangered. The total population in 2008 was less than 600, of which the mature specimens numbered 250. But in 2021, the population has slowly increased from 600 to 1528 individuals. It is confined to four major protected areas: the Senkele Wildlife Sanctuary, Nechisar National Park, Awash National Park and Maze National Park.<ref name=swayne>Template:Cite iucn</ref> The hartebeest in Senkele have to compete with the livestock of the Oromo people.<ref name=lewis/> A study in the Nechisar National Park during 2009 and 2010 found a considerable increase in the livestock of the Oromos (49.9% and 56.5% increase during 2006 and 2010, respectively), illegal resource exploitation, and habitat loss as major threats to the Swayne's hartebeest populations there.<ref name=datiko>Template:Cite journal</ref>
- The western hartebeest is listed as Near Threatened.<ref name=major>Template:Cite iucn</ref> It has been eliminated from most of its range, including the southwestern savannas and Boucle du Baoulé National Park in Mali; southwestern Niger; southern Senegal; Gambia; Ivory Coast; Burkina Faso. Small populations survive in Bafing National Park and the area bounded by Bamako, Bougouni and Sikasso in Mali; Tamou Reserve in Niger; Niokolo-Koba National Park in Senegal; Comoé National Park in Ivory Coast; Diefoula forest and Nazinga Game Ranch in Burkina Faso; Pendjari National Park in Benin; and Bouba Njida, Bénoué, and Faro National Parks in Cameroon.<ref name=east/>
Relationship with humansEdit
Hartebeest are popular game and trophy animals as they are prominently visible and hence easy to hunt.<ref name=kingdon/><ref name=awf/> Pictorial as well as epigraphic evidence from Egypt suggests that in the Upper Palaeolithic age, Egyptians hunted hartebeest and domesticated them. The hartebeest was a prominent source of meat,<ref name="neer">Template:Cite book</ref> but its economic significance was lower than that of gazelles and other desert species.<ref name=heckel/> However, from the beginning of the Neolithic age, hunting became less common and consequently the remains of the hartebeest from this period in ancient Egypt, where it is now extinct, are rare.<ref name=neer/>
In a study on the effect of place and sex on carcass characteristics, the average carcass weight of the male red hartebeest was Template:Convert and that of females was Template:Convert. The meat of the animals from Qua-Qua region had the highest lipid content—Template:Convert per Template:Convert of meat. Negligible differences were found in the concentrations of individual fatty acids, amino acids, and minerals. The study considered hartebeest meat to be healthy, as the ratio of polyunsaturated to saturated fatty acids was 0.78, slightly more than the recommended 0.7.<ref name=hoffman>Template:Cite journal</ref>
ReferencesEdit
External linksEdit
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