The hartebeest (; 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 at the shoulder, and has a typical head-and-body length of . The weight ranges from . 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 slenderer. Horns can reach lengths of . 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.
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 . 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.
Etymology 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, a Swahili word. 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). 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. 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. Vrba dissolved the new genus in 1997 after reconsideration. 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. 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. Another analysis of cytochrome b and D-loop sequence data shows a notable affinity between the A. b. lelwel and A. b. tora divisions.
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. It was declared extinct in 1994 by the International Union for the Conservation of Nature and Natural Resources (IUCN). 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. The population of this hartebeest is on the rise. A. b. cokii Günther, 1884: Known as Coke's hartebeest or kongoni. Native to and confined within Kenya and northern Tanzania. 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. 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. 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. 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. Confirmed as endangered species, its populations are low with at least 1500 individuals, and on verge of population decline.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 A. b. tora Gray, 1873 : Known as the tora hartebeest. Formerly occurred in northwestern Ethiopia and western and southwestern Eritrea. Its present status is unclear, though locals have reported small numbers from these areas.
Genetics and hybrids
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.
The diploid number of chromosomes in the hartebeest is 40. Hybrids are usually reported from areas where ranges of two subspecies overlap. 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. 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.
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. 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. 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. 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.