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The hartebeest (; ''Alcelaphus buselaphus''), also known as kongoni or kaama, is an
African African or Africans may refer to: * Anything from or pertaining to the continent of Africa: ** People who are native to Africa, descendants of natives of Africa, or individuals who trace their ancestry to indigenous inhabitants of Africa *** Ethn ...
antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
. It is the only member of the genus ''Alcelaphus''. Eight
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
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 Horns or The Horns may refer to: * Plural of Horn (instrument), a group of musical instruments all with a horn-shaped bells * The Horns (Colorado), a summit on Cheyenne Mountain * ''Horns'' (novel), a dark fantasy novel written in 2010 by Joe Hill ...
, a short neck, and pointed ears. Its legs, which often have black markings, are unusually long. The
coat A coat typically is an outer garment for the upper body as worn by either gender for warmth or fashion. Coats typically have long sleeves and are open down the front and closing by means of buttons, zippers, hook-and-loop fasteners, toggles, a ...
is generally short and shiny. Coat colour varies by the subspecies, from the sandy brown of the
western hartebeest The western hartebeest (''Alcelaphus buselaphus major'') is an antelope native to the medium to tall grassland plains of Benin, Burkina Faso, Cameroon, the Central African Republic, Chad, Côte d'Ivoire, Ghana, Guinea, Guinea-Bissau, Mali, Niger, ...
to the chocolate brown of the
Swayne's hartebeest Swayne's hartebeest (''Alcelaphus buselaphus swaynei'') is an endangered antelope native to Ethiopia. Two of the largest remaining populations are located in Senkelle Swayne's Hartebeest Sanctuary, Nechisar National Park and Maze National Park. It ...
. Both sexes of all subspecies have horns, with those of females being more slender. 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
herd A herd is a social group of certain animals of the same species, either wild or domestic. The form of collective animal behavior associated with this is called ''herding''. These animals are known as gregarious animals. The term ''herd'' is ...
s 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 Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at one to two years of age.
Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
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
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
s and wooded grasslands, hartebeest often move to more arid places after rainfall. They have been reported from altitudes on
Mount Kenya Mount Kenya (Kikuyu: ''Kĩrĩnyaga'', Kamba, ''Ki Nyaa'') is the highest mountain in Kenya and the second-highest in Africa, after Kilimanjaro. The highest peaks of the mountain are Batian (), Nelion () and Point Lenana (). Mount Kenya is locat ...
up to . The hartebeest was formerly widespread in Africa, but populations have undergone drastic decline due to
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, 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 The bubal hartebeest, also known as northern hartebeest or bubal antelope or simply bubal (''Alcelaphus buselaphus buselaphus'') is the extinct nominal (i.e., first described) subspecies of hartebeest. It was formerly found north of the Saharan De ...
was declared
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN) in 1994. While the populations of the red hartebeest are on the rise, those of the
Tora hartebeest The tora hartebeest, or simply tora (''Alcelaphus buselaphus tora''), is an extremely endangered antelope, native to Eritrea and Ethiopia. It has possibly been extirpated from Sudan. One of the most critically endangered large mammals in the wor ...
, already Critically Endangered, are falling. The hartebeest is extinct in Algeria, Egypt, Lesotho, Libya, Morocco, Somalia, and Tunisia; but has been introduced into
Eswatini Eswatini ( ; ss, eSwatini ), officially the Kingdom of Eswatini and formerly named Swaziland ( ; officially renamed in 2018), is a landlocked country in Southern Africa. It is bordered by Mozambique to its northeast and South Africa to its no ...
and
Zimbabwe Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and Mozam ...
. It is a popular
game animal Game or quarry is any wild animal hunted for animal products (primarily meat), for recreation (" sporting"), or for trophies. The species of animals hunted as game varies in different parts of the world and by different local jurisdictions, thou ...
due to its highly regarded meat.


Etymology

The vernacular name "hartebeest" may have originated from the obsolete
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
word ''hertebeest'', literally ''deer beast'', based on the resemblance (to early
Dutch settlers Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
) of the antelope to
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
. The first use of the word "hartebeest" in South African literature was in Dutch colonial administrator
Jan van Riebeeck Johan Anthoniszoon "Jan" van Riebeeck (21 April 1619 – 18 January 1677) was a Dutch navigator and colonial administrator of the Dutch East India Company. Life Early life Jan van Riebeeck was born in Culemborg, as the son of a surgeon. He ...
's journal ''Daghregister'' in 1660. He wrote: "''Meester Pieter ein hart-beest geschooten hadde'' (Master Pieter an Meerhoffhad 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 In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
,
Coke's hartebeest Coke's hartebeest (''Alcelaphus buselaphus cokii'') or ''Kongoni'' is a large migratory antelope that is native to Kenya and Tanzania. It can breed with Lelwel hartebeest to produce a hybrid known as the Kenya Highland hartebeest (''Alcelaphus ...
.


Taxonomy

The
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
of the hartebeest is ''Alcelaphus buselaphus''. First described by German zoologist
Peter Simon Pallas Peter Simon Pallas Fellow of the Royal Society, FRS FRSE (22 September 1741 – 8 September 1811) was a Prussian zoologist and botanist who worked in Russia between 1767 and 1810. Life and work Peter Simon Pallas was born in Berlin, the son ...
in 1766, it is classified in the genus ''Alcelaphus'' and placed in the family
Bovidae The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, ...
. In 1979, palaeontologist
Elisabeth Vrba Elisabeth S. Vrba (born 17 May 1942) is a paleontologist at Yale University who developed the turnover-pulse hypothesis. Education Vrba earned her Ph.D. in Zoology and Palaeontology at the University of Cape Town, in 1974. Vrba studied zoology ...
supported ''
Sigmoceros Lichtenstein's hartebeest (''Alcelaphus buselaphus lichtensteinii'')Wilson, Don E. & Reeder, DeeAnn M. (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 ppAvailable ...
'' as a separate genus for
Lichtenstein's hartebeest Lichtenstein's hartebeest (''Alcelaphus buselaphus lichtensteinii'')Wilson, Don E. & Reeder, DeeAnn M. (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 ppAvailable ...
, 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 A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
, 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 A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more. ...
, who classified it as an independent species of ''Alcelaphus''. Zoologists such as
Jonathan Kingdon Jonathan Kingdon (born 1935 in Tanzania) is a zoologist, science author, and artist; a research associate at the University of Oxford. He focuses on taxonomic illustration and evolution of the mammals of Africa. He is a contributor to The Oxford B ...
and
Theodor Haltenorth Dr. Theodor Haltenorth (May 18, 1910 – January 30, 1981) was a German mammalogist. He worked mainly in Munich, Bavaria, Germany, and was a key figure in the Quagga Project. He also taxonomised the Cretan wildcat The Cretan wildcat is a me ...
considered it to be a subspecies of ''A. buselaphus''. Vrba dissolved the new genus in 1997 after reconsideration. An
MtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
analysis could find no evidence to support a separate genus for Lichtenstein's hartebeest. It also showed the tribe
Alcelaphini The subfamily Alcelaphinae or tribe Alcelaphini of the family Bovidae contains wildebeest, hartebeest, bonteboks, and several similar species. Depending on the classification, there are 6–10 species placed in four genera, although ''Beatragus' ...
to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, and discovered close affinity between the ''Alcelaphus'' and the sassabies (genus ''
Damaliscus ''Damaliscus'', commonly known as tsessebes, is a genus of antelope in the family Bovidae, subfamily Alcelaphinae The subfamily Alcelaphinae or tribe Alcelaphini of the family Bovidae contains wildebeest, hartebeest, bonteboks, and severa ...
'')—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 The University of Copenhagen ( da, Københavns Universitet, KU) is a prestigious public university, public research university in Copenhagen, Copenhagen, Denmark. Founded in 1479, the University of Copenhagen is the second-oldest university in ...
and colleagues, which sampled the control region of the
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
, found that these two formed a clade within ''A. buselaphus'', and that recognising these as species would render ''A. buselaphus''
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
(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 Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
subspecies. Conversely a 2001
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
study, based on D–loop and
cytochrome b Cytochrome b within both molecular and cell biology, is a protein found in the mitochondria of eukaryotic cells. It functions as part of the electron transport chain and is the main subunit of transmembrane cytochrome bc1 and b6f complexes. F ...
analysis by Øystein Flagstad (of the Norwegian Institute for Nature Research,
Trondheim Trondheim ( , , ; sma, Tråante), historically Kaupangen, Nidaros and Trondhjem (), is a city and municipality in Trøndelag county, Norway. As of 2020, it had a population of 205,332, was the third most populous municipality in Norway, and ...
) 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 Nomination is part of the process of selecting a candidate for either election to a public office, or the bestowing of an honor or award. A collection of nominees narrowed from the full list of candidates is a short list. Political office In the ...
, 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 Affinity may refer to: Commerce, finance and law * Affinity (law), kinship by marriage * Affinity analysis, a market research and business management technique * Affinity Credit Union, a Saskatchewan-based credit union * Affinity Equity Partn ...
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 The bubal hartebeest, also known as northern hartebeest or bubal antelope or simply bubal (''Alcelaphus buselaphus buselaphus'') is the extinct nominal (i.e., first described) subspecies of hartebeest. It was formerly found north of the Saharan De ...
or northern hartebeest. Formerly occurred across northern Africa, from
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
to
Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a transcontinental country spanning the northeast corner of Africa and southwest corner of Asia via a land bridge formed by the Sinai Peninsula. It is bordered by the Mediter ...
. It was exterminated by the 1920s. It was declared extinct in 1994 by the
International Union for the Conservation of Nature and Natural Resources The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
(IUCN). * ''A. b. caama'' ( Saint-Hilaire, 1803) : Known as the
red hartebeest The red hartebeest (''Alcelaphus buselaphus caama''), also called the Cape hartebeest or Caama, is a subspecies of the hartebeest found in Southern Africa. More than 130,000 individuals live in the wild. The red hartebeest is closely related to t ...
or Cape hartebeest. Formerly occurred in southern
Angola , national_anthem = " Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...
; northern and eastern savannahs of
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
; central, southern and southwestern
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label=Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kalahar ...
;
Northern Cape The Northern Cape is the largest and most sparsely populated province of South Africa. It was created in 1994 when the Cape Province was split up. Its capital is Kimberley. It includes the Kalahari Gemsbok National Park, part of the Kgalagadi T ...
,
Eastern Cape The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...
,
Western Cape The Western Cape is a province of South Africa, situated on the south-western coast of the country. It is the fourth largest of the nine provinces with an area of , and the third most populous, with an estimated 7 million inhabitants in 2020 ...
, Free State,
Northwest The points of the compass are a set of horizontal, radially arrayed compass directions (or azimuths) used in navigation and cartography. A compass rose is primarily composed of four cardinal directions—north, east, south, and west—each sep ...
and
Gauteng Gauteng ( ) is one of the nine provinces of South Africa. The name in Sotho-Tswana languages means 'place of gold'. Situated on the Highveld, Gauteng is the smallest province by land area in South Africa. Although Gauteng accounts for only ...
provinces and western
KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN and known as "the garden province") is a province of South Africa that was created in 1994 when the Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu) and Natal Province were merged. It is locate ...
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 Coke's hartebeest (''Alcelaphus buselaphus cokii'') or ''Kongoni'' is a large migratory antelope that is native to Kenya and Tanzania. It can breed with Lelwel hartebeest to produce a hybrid known as the Kenya Highland hartebeest (''Alcelaphus ...
or kongoni. Native to and confined within
Kenya ) , national_anthem = "Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
and northern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
. * ''A. b. lelwel'' (Heuglin, 1877) : Known as the
Lelwel hartebeest The Lelwel hartebeest (''Alcelaphus buselaphus lelwel''), also known as Jackson's hartebeest, is an antelope native to Central African Republic, Chad, the Democratic Republic of the Congo, Ethiopia, Kenya, Sudan, Tanzania, and Uganda. The Lel ...
. Formerly found in northern and northeastern
Democratic Republic of the Congo The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
; southeastern and southwestern
Sudan Sudan ( or ; ar, السودان, as-Sūdān, officially the Republic of the Sudan ( ar, جمهورية السودان, link=no, Jumhūriyyat as-Sūdān), is a country in Northeast Africa. It shares borders with the Central African Republic t ...
; and the northwestern extreme of
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
. Drastic population decrease since the 1980s has confined most individuals to protected areas inside and outside its range. * ''A. b. lichtensteinii'' (
Peters Peters may refer to: People * Peters (surname) * Peters Band, a First Nations band in British Columbia, Canada Places United States * Peters, California, a census-designated place * Peters, Florida, a town * Peters Township, Kingman County, Kan ...
, 1849)
: Known as
Lichtenstein's hartebeest Lichtenstein's hartebeest (''Alcelaphus buselaphus lichtensteinii'')Wilson, Don E. & Reeder, DeeAnn M. (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 ppAvailable ...
. Inhabits the
miombo woodland The Miombo woodland is a tropical and subtropical grasslands, savannas, and shrublands biome (in the World Wide Fund for Nature scheme) located primarily in Central Africa. It includes four woodland savanna ecoregions (listed below) characterized b ...
s of eastern and southern Africa. It is native to Angola, the Democratic Republic of Congo,
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northeast ...
,
Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi ...
, South Africa, Tanzania,
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most cent ...
and
Zimbabwe Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and Mozam ...
. * ''A. b. major'' ( Blyth, 1869) : Known as the
western hartebeest The western hartebeest (''Alcelaphus buselaphus major'') is an antelope native to the medium to tall grassland plains of Benin, Burkina Faso, Cameroon, the Central African Republic, Chad, Côte d'Ivoire, Ghana, Guinea, Guinea-Bissau, Mali, Niger, ...
. Formerly occurred widely in Mali,
Niger ) , official_languages = , languages_type = National languagesSenegal Senegal,; Wolof: ''Senegaal''; Pulaar: 𞤅𞤫𞤲𞤫𞤺𞤢𞥄𞤤𞤭 (Senegaali); Arabic: السنغال ''As-Sinighal'') officially the Republic of Senegal,; Wolof: ''Réewum Senegaal''; Pulaar : 𞤈𞤫𞤲𞤣𞤢𞥄𞤲𞤣𞤭 ...
,
Gambia The Gambia,, ff, Gammbi, ar, غامبيا officially the Republic of The Gambia, is a country in West Africa. It is the smallest country within mainland AfricaHoare, Ben. (2002) ''The Kingfisher A-Z Encyclopedia'', Kingfisher Publicatio ...
,
Guinea-Bissau Guinea-Bissau ( ; pt, Guiné-Bissau; ff, italic=no, 𞤘𞤭𞤲𞤫 𞤄𞤭𞤧𞤢𞥄𞤱𞤮, Gine-Bisaawo, script=Adlm; Mandinka: ''Gine-Bisawo''), officially the Republic of Guinea-Bissau ( pt, República da Guiné-Bissau, links=no ) ...
,
Guinea Guinea ( ),, fuf, 𞤘𞤭𞤲𞤫, italic=no, Gine, wo, Gine, nqo, ߖߌ߬ߣߍ߫, bm, Gine officially the Republic of Guinea (french: République de Guinée), is a coastal country in West Africa. It borders the Atlantic Ocean to the we ...
,
Ivory Coast Ivory Coast, also known as Côte d'Ivoire, officially the Republic of Côte d'Ivoire, is a country on the southern coast of West Africa. Its capital is Yamoussoukro, in the centre of the country, while its largest city and economic centre is ...
,
Ghana Ghana (; tw, Gaana, ee, Gana), officially the Republic of Ghana, is a country in West Africa. It abuts the Gulf of Guinea and the Atlantic Ocean to the south, sharing borders with Ivory Coast in the west, Burkina Faso in the north, and To ...
,
Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...
, southwestern Chad, Cameroon, western Central African Republic and
Benin Benin ( , ; french: Bénin , ff, Benen), officially the Republic of Benin (french: République du Bénin), and formerly Dahomey, is a country in West Africa. It is bordered by Togo to the west, Nigeria to the east, Burkina Faso to the north ...
. Nowadays it occurs in much lower numbers mainly in protected areas of these countries. It is probably extinct in Gambia. * ''A. b. swaynei'' (
Sclater Philip Lutley Sclater (4 November 1829 – 27 June 1913) was an English lawyer and zoologist. In zoology, he was an expert ornithologist, and identified the main zoogeographic regions of the world. He was Secretary of the Zoological Society o ...
, 1892)
: Known as
Swayne's hartebeest Swayne's hartebeest (''Alcelaphus buselaphus swaynei'') is an endangered antelope native to Ethiopia. Two of the largest remaining populations are located in Senkelle Swayne's Hartebeest Sanctuary, Nechisar National Park and Maze National Park. It ...
. Restricted to the southern
Rift Valley A rift valley is a linear shaped lowland between several highlands or mountain ranges created by the action of a geologic rift. Rifts are formed as a result of the pulling apart of the lithosphere due to extensional tectonics. The linear dep ...
in Ethiopia. It formerly occurred throughout the Rift Valley, and its range extended eastward into northwestern
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constituti ...
. It has disappeared from Somalia by 1930. Its populations are very low and on the decline. * ''A. b. tora''
Gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed o ...
, 1873
: Known as the
tora hartebeest The tora hartebeest, or simply tora (''Alcelaphus buselaphus tora''), is an extremely endangered antelope, native to Eritrea and Ethiopia. It has possibly been extirpated from Sudan. One of the most critically endangered large mammals in the wor ...
. Formerly occurred in northwestern Ethiopia and western and southwestern
Eritrea Eritrea ( ; ti, ኤርትራ, Ertra, ; ar, إرتريا, ʾIritriyā), officially the State of Eritrea, is a country in the Horn of Africa region of Eastern Africa, with its capital and largest city at Asmara. It is bordered by Ethiopia ...
. 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 Senkelle Swayne's Hartebeest Sanctuary is a protected area in the Oromia Region (or ''kilil'') of Ethiopia, dedicated especially to the protection of the Swayne's hartebeest (''Alcelaphus buselaphus swaynei''). Covering 54 square kilometers, the ...
and the
Nechisar National Park Nechisar National Park (or Nech Sar National Park) is a national park in the Southern Nations, Nationalities, and Peoples' Region (SNNPR) of Ethiopia. It is in the Great Rift Valley, within the southwestern Ethiopian Highlands. Geography The pa ...
, for mitochondrial (D-loop) and nuclear (
microsatellite A microsatellite is a tract of repetitive DNA in which certain DNA motifs (ranging in length from one to six or more base pairs) are repeated, typically 5–50 times. Microsatellites occur at thousands of locations within an organism's genome. ...
) variability in an attempt to estimate the levels of
genetic variation Genetic variation is the difference in DNA among individuals or the differences between populations. The multiple sources of genetic variation include mutation and genetic recombination. Mutations are the ultimate sources 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 Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...
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 ''In-situ'' conservation is the on-site conservation or the conservation of genetic resources in natural populations of plant or animal species, such as forest genetic resources in natural populations of Teagan species. This process protects th ...
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 Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
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 The Blue Nile (; ) is a river originating at Lake Tana in Ethiopia. It travels for approximately through Ethiopia and Sudan. Along with the White Nile, it is one of the two major tributaries of the Nile and supplies about 85.6% of the water ...
to about 9° N latitude. A study proved a male hybrid of the red hartebeest and the
blesbok The blesbok or blesbuck (''Damaliscus pygargus phillipsi'') is a subspecies of the bontebok antelope endemic to South Africa, Eswatini and Namibia. It has a distinctive white face and forehead which inspired the name, because ''bles'' is the Afri ...
(''Damaliscus pygargus'') to be
sterile Sterile or sterility may refer to: *Asepsis, a state of being free from biological contaminants * Sterile (archaeology), a sediment deposit which contains no evidence of human activity *Sterilization (microbiology), any process that eliminates or ...
. Sterility of the hybrid was attributed to difficulties in
segregation Segregation may refer to: Separation of people * Geographical segregation, rates of two or more populations which are not homogenous throughout a defined space * School segregation * Housing segregation * Racial segregation, separation of humans ...
during
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
, indicated by
azoospermia Azoospermia is the medical condition of a man whose semen contains no sperm. It is associated with male infertility, but many forms are amenable to medical treatment. In humans, azoospermia affects about 1% of the male population and may be seen ...
and a low number of
germ cell Germ or germs may refer to: Science * Germ (microorganism), an informal word for a pathogen * Germ cell, cell that gives rise to the gametes of an organism that reproduces sexually * Germ layer, a primary layer of cells that forms during embry ...
s in its
seminiferous tubule Seminiferous tubules are located within the testes, and are the specific location of meiosis, and the subsequent creation of male gametes, namely spermatozoa. Structure The epithelium of the tubule consists of a type of sustentacular cells known ...
s. 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 Lake Victoria is one of the African Great Lakes. With a surface area of approximately , Lake Victoria is Africa's largest lake by area, the world's largest tropical lake, and the world's second-largest fresh water lake by surface area after ...
and
Mount Kenya Mount Kenya (Kikuyu: ''Kĩrĩnyaga'', Kamba, ''Ki Nyaa'') is the highest mountain in Kenya and the second-highest in Africa, after Kilimanjaro. The highest peaks of the mountain are Batian (), Nelion () and Point Lenana (). Mount Kenya is locat ...
, but is now believed to be restricted to the Lambwe Valley (south-west Kenya) and
Laikipia Laikipia County is one of the 47 Counties of Kenya, located on the Equator in the former Rift Valley Province of the Country. Laikipia is a cosmopolitan County and is Listed as County number 31. The county has two major urban centres: Nanyuki t ...
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. * ''Alcelaphus lelwel x swaynei'' : Also known as the Neumann's hartebeest, named after traveller and hunter
Arthur Henry Neumann Arthur Henry Neumann (12 June 1850 – 29 May 1907) was an English explorer, hunter, soldier, farmer and travel writer famous for his exploits in Equatorial East Africa. In 1898 he published ''Elephant Hunting in East Equatorial Africa''.A H Neum ...
. 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 The Omo River (also called Omo-Bottego) in southern Ethiopia is the largest Ethiopian river outside the Nile Basin. Its course is entirely contained within the boundaries of Ethiopia, and it empties into Lake Turkana on the border with Kenya. The ...
and north of
Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern end crossing into Ethiopia. It is the world's largest permanent desert lake and the world's largest alkaline lake. B ...
, stretching north-east of
Lake Chew Bahir Lake Chew Bahir (Amharic: ጨው ባሕር ''č̣ew bāhir'', "salty lake") or Lake Istifanos, also called Stefanie, Basso Naebor and Chuwaha, is a lake in southern Ethiopia, located on the southwestern end of the Southern Nations, Nationalities ...
to near
Lake Chamo Lake Chamo ( Amharic: ቻሞ ሐይቅ) is a lake in the Southern Nations, Nationalities, and Peoples' Region of southern Ethiopia. Located in the Main Ethiopian Rift, it is at an elevation of 1,110 meters. The Chamo lake is just to the south of ...
.


Evolution

The genus ''Alcelaphus'' emerged about 4.4 million years ago in a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
whose other members were ''
Damalops ''Damalops'' is an extinct genus of Alcelaphinae. It lived during the Pliocene and Pleistocene in southern Asia, where the species ''Damalops palaeindicus'' is known from the Siwaliks in the northern part of the India India, officially th ...
'', ''
Numidocapra ''Numidocapra'' is an extinct genus of bovid from the Pleistocene of Africa. Taxonomy Originally placed in the subfamily caprinae, ''Numidocapra'' is now generally classified as a member of the subfamily Alcelaphinae. Three species are considere ...
'', '' Rabaticeras'', ''
Megalotragus ''Megalotragus'' was a genus of very large extinct African alcelaphines that occurred from the Pliocene to early Holocene.Thackeray, John Francis. (2015). Faunal Remains from Holocene Deposits, Excavation 1, Wonderwerk Cave, South Africa. Africa ...
'', '' Oreonagor'', and ''Connochaetes''. An analysis using
phylogeographic Phylogeography is the study of the historical processes that may be responsible for the past to present geographic distributions of genealogical lineages. This is accomplished by considering the geographic distribution of individuals in light of ge ...
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 The hirola (''Beatragus hunteri''), also called the Hunter's hartebeest or Hunter's antelope, is a critically endangered antelope species found on the border between Kenya and Somalia. It was first described by the big game hunter and zoologist ...
). 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 A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
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. The earliest fossil record dates back to nearly 0.7 million years ago. Fossils of the
red hartebeest The red hartebeest (''Alcelaphus buselaphus caama''), also called the Cape hartebeest or Caama, is a subspecies of the hartebeest found in Southern Africa. More than 130,000 individuals live in the wild. The red hartebeest is closely related to t ...
have been found in Elandsfontein, Cornelia (Free State) and
Florisbad Florisbad is a health resort 45 km northwest of Bloemfontein and 47 km south-west of Brandfort, near the Haagenstad salt-pan. Named after Floris Venter who opened up the mineral spring. Florisbad archaeological and paleontological site ...
in
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
, as well as in
Kabwe Kabwe is the capital of the Zambian Central Province and the Kabwe District, with a population estimated at 202,914 at the 2010 census. Named Broken Hill until 1966, it was founded when lead and zinc deposits were discovered in 1902. Kabwe also ...
in
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most cent ...
. In
Israel Israel (; he, יִשְׂרָאֵל, ; ar, إِسْرَائِيل, ), officially the State of Israel ( he, מְדִינַת יִשְׂרָאֵל, label=none, translit=Medīnat Yīsrāʾēl; ), is a country in Western Asia. It is situated ...
, hartebeest remains have been found in northern
Negev The Negev or Negeb (; he, הַנֶּגֶב, hanNegév; ar, ٱلنَّقَب, an-Naqab) is a desert and semidesert region of southern Israel. The region's largest city and administrative capital is Beersheba (pop. ), in the north. At its southe ...
,
Shephelah The Shephelah or Shfela, lit. "lowlands" ( hbo, הַשְּפֵלָה ''hašŠǝfēlā'', also Modern Hebrew: , ''Šǝfēlat Yəhūda'', the "Judaean foothills"), is a transitional region of soft-sloping rolling hills in south-central Israel str ...
,
Sharon Plain The Sharon plain ( ''HaSharon Arabic: سهل شارون Sahel Sharon'') is the central section of the Israeli coastal plain. The plain lies between the Mediterranean Sea to the west and the Samarian Hills, to the east. It stretches from Nahal ...
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 primary production, 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 western hartebeest is a pale sandy-brown, but the front of the legs are darker. 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. The Lelwel hartebeest is a reddish tan. Coke's hartebeest is reddish to tawny in the upper parts, but has relatively lighter legs and rump. Lichtenstein's hartebeest is reddish brown, though the flanks are a lighter tan and the rump whitish. 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. 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. Fine textured, the body hair of the hartebeest is about long. 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. Both sexes of all subspecies have horns, with those of females being more slender. Horns can reach lengths of ; the maximum horn length is , recorded from a Namibian red hartebeest. 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. 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. 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. The horns of Swayne's hartebeest are thin and shaped like parentheses, curving upward and then backward. The horns of the Tora hartebeest are particularly thin and spread out sideways, diverging more than in any other subspecies. Apart from its long face, the large chest and the sharply sloping back differentiate the hartebeest from other antelopes. The hartebeest shares several physical traits with the sassabies (genus ''Damaliscus''), such as an elongated and narrow face, the shape of the horns, the coat (animal), pelage texture and colour, and the terminal tuft of the tail. The wildebeest have more specialised skull and horn features than the hartebeest. 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. 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. However, moving herds are not so cohesive, and tend to disperse frequently. The members of a herd can be divided into four groups: Territory (animal), 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. Sparring between males and females is common. 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. The male marks the border of his territory through defecation. Hartebeest are remarkably alert and cautious animals with highly developed brains. 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. Adult hartebeest are preyed upon by lions, leopards, hyenas and Lycaon pictus, wild dogs; cheetahs and jackals target juvenile (organism), juveniles. Crocodiles may also prey on hartebeest. 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 horns are also used during fights among males for Dominance (ethology), dominance in the breeding season; the clash of the horns is loud enough that it can be heard from hundreds of metres away. 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. Fights are rarely serious, but can be fatal if they are. 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. The hartebeest uses defecation as an olfactory and visual display. Herds are generally sedentary, and tend to migrate only under adverse conditions such as natural calamities. The hartebeest is the least migratory in the Tribe (biology), 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.


Parasites and diseases

Several parasites have been isolated from the hartebeest. These parasites regularly alternate between hartebeest and gazelles or wildebeest. Hartebeest can be infected with East coast fever, theileriosis due to ''Rhipicephalus evertsi'' and ''Theileria'' species. South of the Sahara, common parasites include ''Loewioestrus variolosus'', ''Gedoelstia cristata'' and ''Gedoelstia hassleri, G. hassleri''. The latter two species can cause serious diseases such as encephalitis. However, parasites are not always harmful – 252 Larva (zoology), larvae were found in the head of one Zambian individual without any pathogenicity. Nematodes, cestodes, Paramphistomum, 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. The hartebeest has much lower food intake than the other members of Alcelaphini. The long thin Muzzle (animal), 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. 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. These unique abilities could have allowed the hartebeest to prevail over other animals millions of years ago, leading to its successful radiation across Africa. Grasses generally comprise at least 80 percent of the hartebeest's diet, but they account for over 95 percent 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 Culm (botany), culms of grasses. 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. 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.


Reproduction

Mating in hartebeest takes place throughout the year, with one or two peaks that can be influenced by the availability of food. Both males and females reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...
at one to two years of age. Reproduction varies by the subspecies and local factors. Mating takes place in the territories defended by a single male, mostly in open areas. The males may fight fiercely for dominance, following which the dominant male smells the female's genitalia, and follows her if she is in Estrous cycle, oestrus. Sometimes a female in oestrus holds out her tail slightly to signal her receptivity, 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. Any intruder at this time is chased away. In large herds, females often mate with several males.
Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
is eight to nine months long, after which a single calf weighing about 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. Though calves can move about on their own shortly after birth, they usually lie in the open in close proximity of their mothers. The calf is weaned at four months, but young males stay with their mothers for two and a half years, longer than in other Alcelaphini. 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, 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. They have been reported from altitudes on
Mount Kenya Mount Kenya (Kikuyu: ''Kĩrĩnyaga'', Kamba, ''Ki Nyaa'') is the highest mountain in Kenya and the second-highest in Africa, after Kilimanjaro. The highest peaks of the mountain are Batian (), Nelion () and Point Lenana (). Mount Kenya is locat ...
up to . The red hartebeest is known to move across large areas, and females roam home ranges of over , with male territories in size. 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 International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. The species as a whole is classified as Least Concern by the IUCN. The hartebeest is extinct in Algeria, Egypt, Lesotho, Libya, Morocco, Somalia, and Tunisia. * The Bubal hartebeest has been declared extinct since 1994. German explorer Heinrich Barth, in his works of 1857, cites firearms and European intrusion among the reasons for the decrease in its numbers. 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 Region, 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. Fewer than 70,000 individuals are left. Most of the population nowadays is found in Chad in the Salamat Region, 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. *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. * 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. Its population is estimated to be over 130,000 (as of 2008), mostly in southern Africa. In Namibia, the largest population occurs in the Etosha National Park. A reintroduced population is flourishing in the Malolotja Nature Reserve (
Eswatini Eswatini ( ; ss, eSwatini ), officially the Kingdom of Eswatini and formerly named Swaziland ( ; officially renamed in 2018), is a landlocked country in Southern Africa. It is bordered by Mozambique to its northeast and South Africa to its no ...
), outside its range. However, numbers have seen a sharp fall in southwestern Botswana. *The Tora hartebeest is listed as Critically Endangered; the IUCN has ascertained that fewer than 250 mature individuals survive as of 2008. They are possibly extinct in Sudan due to excessive hunting and agricultural expansion, but may still exist in smaller numbers in Eritrea and Ethiopia. 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. *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 within 250. It is confined to four major protected areas: the
Senkele Wildlife Sanctuary Senkelle Swayne's Hartebeest Sanctuary is a protected area in the Oromia Region (or ''kilil'') of Ethiopia, dedicated especially to the protection of the Swayne's hartebeest (''Alcelaphus buselaphus swaynei''). Covering 54 square kilometers, the ...
,
Nechisar National Park Nechisar National Park (or Nech Sar National Park) is a national park in the Southern Nations, Nationalities, and Peoples' Region (SNNPR) of Ethiopia. It is in the Great Rift Valley, within the southwestern Ethiopian Highlands. Geography The pa ...
, Awash National Park and Maze National Park. The hartebeest in Senkele have to compete with the livestock of the Oromo people. 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. *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 Ivory Coast, also known as Côte d'Ivoire, officially the Republic of Côte d'Ivoire, is a country on the southern coast of West Africa. Its capital is Yamoussoukro, in the centre of the country, while its largest city and economic centre is ...
; 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 National Park, Bouba Njida, Bénoué National Park, Bénoué, and Faro National Park, Faro National Parks in Cameroon.


Relationship with humans

Hartebeest are popular game animal, game and Trophy hunting, trophy animals as they are prominently visible and hence easy to hunt. Pictorial as well as epigraphy, epigraphic evidence from Prehistoric Egypt, Egypt suggests that in the Upper Palaeolithic, Upper Palaeolithic age, Egyptians hunted hartebeest and domesticated them. The hartebeest was a prominent source of meat, but its economic significance was lower than that of gazelles and other desert species. 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. In a study on the effect of place and sex on carcass characteristics, the average carcass weight of the male red hartebeest was and that of females was . The meat of the animals from Qua-Qua region had the highest lipid content— per of meat. Negligible differences were found in the concentrations of individual fatty acids, amino acids, and dietary mineral, minerals. The study considered hartebeest meat to be healthy, as the ratio of Polyunsaturated fatty acid, polyunsaturated to saturated fatty acids was 0.78, slightly more than the recommended 0.7.


References


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* * {{Authority control Alcelaphus Mammals described in 1766 Mammals of Africa Taxa named by Peter Simon Pallas