Hartebeest

The vernacular name "hartebeest" may have originated from the obsolete Dutch word hertebeest, literally deer beast, based on the resemblance (to early Dutch settlers) of the antelope to deer. 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)". Another name for the hartebeest is kongoni, Kongoni is often used to refer in particular to one of its subspecies, Coke's hartebeest. ==Taxonomy==

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. 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). However, this finding was not replicated by Alan W. Gentry of the Natural History Museum, who classified it as an independent species of Alcelaphus. Zoologists such as Jonathan Kingdon and Theodor Haltenorth considered it to be a subspecies of A. buselaphus. 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. Subspecies 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. 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. The eight subspecies, including the two controversial ones, are: • † 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. • 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. • A. b. cokii Günther, 1884: Known as Coke's hartebeest or kongoni. Native to and confined within Kenya and northern Tanzania. Drastic population decrease since the 1980s has confined most individuals to protected areas inside and outside its range. • A. b. lichtensteinii (Peters, 1849) : Known as Lichtenstein's hartebeest. Inhabits the miombo woodlands of eastern and southern Africa. It is native to Angola, the Democratic Republic of Congo, Malawi, Mozambique, South Africa, Tanzania, Zambia and Zimbabwe. • A. b. tora Gray, 1873 : Known as the tora hartebeest. Formerly occurred in northwestern Ethiopia and western and southwestern Eritrea. 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. • 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. This is considered to be a cross between the Lelwel hartebeest and Swayne's hartebeest. 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. ==Evolution==

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. 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 400,000 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 200,000 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. 700,000 years ago. 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. 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. ==Description==

A large antelope with a particularly elongated forehead and oddly shaped horns, the hartebeest stands just over at the shoulder, and has a typical head-and-body length of . The weight ranges from . The tail, long, ends in a black tuft. The other distinctive features of the hartebeest are its long legs (often with black markings), short neck, and pointed ears. A study correlated the size of hartebeest species to habitat productivity and rainfall. The western hartebeest is the largest subspecies, and has a characteristic white line between the eyes. The red hartebeest is also large, with a black forehead and a contrasting light band between the eyes. The large Lelwel hartebeest has dark stripes on the front of its legs. Coke's hartebeest is moderately large, with a shorter forehead and longer tail in comparison to the other subspecies. Lichtenstein's hartebeest is smaller, with dark stripes on the front of the legs, as in the Lelwel hartebeest. The Swayne's hartebeest is smaller than the Tora hartebeest, but both have a shorter forehead and similar appearance. Generally short and shiny, the coat varies in colour according to subspecies. The Lelwel hartebeest is a reddish tan. 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. 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. ==Ecology and behaviour==

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. 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. 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. 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. Hartebeest can be infected with theileriosis due to Rhipicephalus evertsi and Theileria species. 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. However, parasites are not always harmful – 252 larvae were found in the head of one Zambian individual without any pathogenicity. Nematodes, cestodes, paramphistomes; and the roundworm Setaria labiatopapillosa have also been isolated from the hartebeest. 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. Diet Hartebeest are primarily grazers, and their diets consist mostly of grasses. 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. A study found that the hartebeest is able to digest a higher proportion of food than the topi and the wildebeest. In areas with scarce water, it can survive on melons, roots, and tubers. Reproduction Mating in hartebeest takes place throughout the year, with one or two peaks that can be influenced by the availability of food. 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. The lifespan is 12 to 15 years. ==Habitat==

Hartebeest inhabit dry savannas, open plains and wooded grasslands, Females in the Nairobi National Park (Kenya) have individual home ranges stretching over , which are not particularly associated with any one female group. Average female home ranges are large enough to include 20 to 30 male territories. ==Status and conservation==

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. It was extinct in Tunisia by the late 19th century. The last individual was shot in Missour (Algeria) in 1925. • '''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. • 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. • 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. 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. The hartebeest in Senkele have to compete with the livestock of the Oromo people. • The western hartebeest is listed as Near Threatened. 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. ==Relationship with humans==

Hartebeest are popular game and trophy animals as they are prominently visible and hence easy to hunt. but its economic significance was lower than that of gazelles and other desert species. ==References==