Lesser kudu (Ammelaphus or Tragelaphus imberbis)
The kudu is 1.10-1.75 m from the tip of the snout to the base of the tail and stands 0.9-1.1 m (35-43 in) tall at the shoulder. The tail ranges from 26-30 cm with up to 90 cm of additional hair length. range Females weigh 50-70 kg (110-150 lb) and males weigh 60-105 kg (130–230 lb). Males have horns ranging from 48-91 cm, have two longitudinal keels, twist 2.5 times, and a basal circumference of 15.6-17.1 cm. At the base, they span 6-12 mm; at their ends, they span 25-35 cm. The skull is long with a short cranium. The long nasal bones are narrowed in the center and form a 'V' shaped suture where they meet the frontals. The supraorbital foramina are within indentations and are elongated horizontally. The paraoccipital proccess are flat and wide and the teeth are hypsodont. The kudu occupies semi-arid areas, such as dry, flat, and densely thicketed areas of subtropical and tropical dry shrubland as well as woodlands and hilly land. It is rarely seen in open or cleared areas or long grass (6). It lives in eastern and north-eastern Africa - the Somali-Masai Arid Zone of Ethiopia, Somalia, Kenya, Uganda and Tanzania. Its range extends from ca. 12°N in the Awash area of Ethiopia southward through southern Ethiopia, much of Somalia except the north and northeast (east of 46° E and north of 08°N), most of Kenya except the southwest, extreme southeast Sudan, extreme northeast Uganda to northeast and central Tanzania. It is extinct in Djibouti (6) and its purported former occurrence in Saudi Arabia (7) is based on two sets of horns said to have originated from Saudi Arabia and one from southern Yemen. No live animals have been reported from the area and the true origin of those specimens remains in doubt; they may have been introduced or released from a collection. The kudu is closely associated with Acacia-Commiphora thornbush in north-eastern Africa (6). It has been recorded at about 1,740 m near Mt Kilimanjaro (8). The kudu is fairly solitary and shy. Females usually live in groups of two to five, ranging up to 24; males are often found alone. The kudu is mainly nocturnal and crepuscular and camouflages well when hiding in dense thickets and dry bush after sunrise. Its large ears aid in a well developed sense of hearing that warns it of potential predators. When startled, it will bark and runs in bounding leaps, holding the tail upright to reveal the white underside. It can jump more than 6 m (20 ft) and 2 m (6.6 ft) high and can reach running speeds of around 70 km/h (43 mph). It feeds at dusk and dawn. It is mainly a browser and eats a diverse variety of bush and tree leaves, shoots, twigs, fruits, grasses and herbs. It obtains most of its water requirements from its food plants. When males are large enough, they fight by locking horns and pressing their heads and horns together and try to force their horns down onto the nape of their opponent. Males and females fight each other for superiority, by standing up on hind legs to try to knock each other down, but the larger males normally win. Males show restraint and are never aggressive towards females, but females may butt their heads against the males. When males mount, they lay their neck and head down and onto the females back (11). Each female has her own, independent estrus cycle and is anestrus for only a couple of weeks (9). She is pregnant for 7.4-8.5 months and separates from her small group to give birth to one calf, weighing 4.0-7.5 kg. 50% of the calves die within the first 6 months, from disease and predation by leopards, hunting dogs and spotted hyenas, and only 25% survive to reach 3 years of age. Males begin horn growth after the first 6-9 months and reach full length after 3 years. Young males stay with their mothers for 1.5-2 years, before they leave and travel alone or in small sporadic male groups (10). Young females form small groups with their mother or siblings. The kudu becomes sexually mature at 1.25-1.50 years, but males do not gain social status to reproduce until they reach the age of 4-5 years (11). The average life span is 10-15 years in the wild and 15 years in captive, with one captive reaching 19.8 years (9). IUCN list the kudu as conservation dependent (9), but Lower Risk (1). The total population is estimated to be at least 118,000 (6), about 33% being in protected areas. Numbers are considered to be in decline overall, due to hunting for sport and its meat and horns, habitat loss, overgrazing, increase in pastoralism and outbreaks of rinderpest, which led to a decline in the mid 1990s (6). The horns are hollowed out and used as wind instruments, honey containers and in spiritual rituals as they are thought to house powerful spirits as well as being a symbol of male virility (2,4). In Tsavo National Park, elephant populations have altered the vegetative landscape.The level of decline is predicted to reach at least 25% over a period of three generations (21-24 years), so approaching the threshold for Vulnerable. The kudu will probably persist in the arid scrublands of northeastern Africa, if human and livestock densities stay relatively low in extensive parts of its range such as northern Kenya and southern Ethiopia. Its status may eventually decline to threatened. Its shyness and preference for thick cover help it withstand considerable hunting pressure (6). Lesser kudu are also vulnerable to the rinderpest virus which periodically breaks out and reduces populations (5). The kudu’s long-term survival prospects would be enhanced by improved protection and management of the protected areas that support substantial populations. Its value as a trophy animal gives it high potential for increased revenue generation in extensive bushlands (6).The lesser kudu is part of the American Zoo and Aquarium Association's studbook program which ensures the most possible genetic diversity within the captive breeding population (6). The lesser kudu was placed in the genus Tragelaphus. It was described by Edward Blyth in 1869. It was thought to be a smaller version of the greater kudu, but is now is considered to be a more primitive species, being the most primitive spiral-horned antelope. Its evolutionary line diverged in the late Miocene, possibly 10 million years ago. There may have been an early hybridization between the proto-lesser kudu and proto-nyala, but these lines have been separate for most of the evolutionary history. The lesser kudu being the most basal member of the 'Tragelaphus' group. In 1912, Edmund Heller established the genus Ammelaphus for the Lesser Kudu, the type species being A. strepsiceros (2). The lesser kudu was raised to a genus level by Peter Grubb and Colin Groves in 2011 (3), as it represents an evolutionary line that has remained separate since the end of the Miocene (5.8 million years ago). Grubb and Groves state that Ammelaphus has two species, the northern (A. imberbis from Ethiopia and Somalia) and southern (A. australis from Kenya, Tanzania, Sudan and southern Somalia). The lesser kudu has 38 chromosomes, in both sexes. Unlike other tragelaphids, the X chromosome and Y chromosome are compound and fused with autosomes from ancestors having a greater chromosome number (5).
- 1. IUCN SSC Antelope Specialist Group (2008). "Tragelaphus imberbis". IUCN Red List of Threatened Species. Version 2012.1. International Union for Conservation of Nature. Retrieved 23 July 2012.
- 2. Heller, Edmund (November 2, 1912). New Genera and Races of African Ungulates. Washington D. C.: Smitsonian Institution. p. 15.
- 3. Groves, Colin, Peter Grubb (2011). Ungulate Taxonomy. Baltimore, Md.: Johns Hopkins University Press. p. 139. ISBN 1-4214-0093-6.
- 4. Miller, Melissa, Tim Wild, Steve Shurter. "Lesser kudu".
- 5. Benirschke, K.; Rüedi, D.; Müller, H.; Kumamoto, A.T.; Wagner, K.L.; Downes, H.S. (1 January 1980). "The unusual karyotype of the lesser kudu, Tragelaphus imberbi's". Cytogenetic and Genome Research 26 (2-4): 85–92. doi:10.1159/000131429.
- 6. East, R. (1999). African Antelope Database 1998. IUCN, Gland, Switzerland.
- 7. Harrison, D., P. Bates (1991). The Mammals of Arabia. Kent, England: Harrison Zoological Museum Publications.
- 8. Grimshaw, J.M., N.J. Cordeiro and C.A.H. Foley. (1995). The mammals of Kilimanjaro. Journal of East African Natural History 84:105-139.
- 9. Weigl, R. (2005). Longevity of Mammals in Captivity; From the Living Collections of the World. [Kleine Senckenberg-Reihe 48]. E. Schweizerbart’sche Verlagsbuchhandlung (Nägele und Obermiller), Stuttgart.
- 10. Nowak, R. (1999). Walker's Mammals of the World. Balitimore: The John Hopkins University.
- 11. Walther, F. (1990). Spiral-Horned Antelopes. Pp. 344-359 in S Parker, ed. Grzimek's Encyclopedia of Mammals. New York: McGraw-Hill Publishing Company.
The range of Tragelaphus imberbis, the Lesser Kudu, is restricted to northeast Africa. It was once believed to inhabit Saudi Arabia, however, only two sets of horns have been found that substantiate this claim. BR
(Harrison and Bates 1991; Nowak 1999; Roosevelt and Heller 1914; Walther 1990)
Biogeographic Regions: ethiopian (Native )
The purported former occurrence of this species in Saudi Arabia (Harrison and Bates 1991) is based on two sets of horns said to have originated from Arabia and one from southern Yemen. No live animals have ever been reported from the area, and the true origin of those specimens remains in doubt.
The Lesser Kudu is the largest bovid in Arabia, with the exception of the Oryx (Harrison and Bates 1991). It is 1.10-1.75 meters from the tip of the snout to the base of the tail and stands at a height of 0.90-1.05 meters. The tail ranges from 0.26 to 0.30 meters with up to 0.90 meters of additional hair length. Males have horns that range from 0.48 to 0.91 meters, have two longitudinal keels, twist 2.5 times, and a basal circumference of 156-171 mm. At the base, they span 6-12 mm; at their ends, they span 0.25-0.35 meters. Males have various shades of grey fur, and females are distinctly more reddish brown. One long white stripe runs the length of the back with 11-14 stripes branching downward off of it. The face has a black stripe from each eye to the nose and a white stripe from each eye to the center of the face. The lip area is white and four white spots are found on the lower jaw (two per side). The legs are fawn colored with a white patch above the hoof. The tail is brown above and white underneath, and a black tip. There are white patches on the throat and chest with a black stripe that spans the chest area. The abdomen is pure white. Young Kudus are similar in color to females but redder, and males become more grey by the age of two years.
The lesser kudu is hard to observe in dry bush due to its camouflage, and because its large ears aid in a well developed sense of hearing that warns it of potential predators.
The skull of the Lesser Kudu is long with a short cranium. The nasal bones are long and narrowed in the center. They form a 'V' shaped suture where they meet the frontals. The supraorbital foramina are located within indentations and are elongated horizontally. The paraoccipital proccess are flat and wide, and the teeth are hypsodont.
(Harrison and Bates 1991; Nowak 1999; Roosevelt and Heller 1914; Walther 1990)
Range mass: 60 to 105 kg.
Other Physical Features: endothermic ; bilateral symmetry
Catalog Number: USNM 182073
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Preparation: Skin; Skull
Collector(s): M. Johnston
Year Collected: 1911
Locality: Longaya Water, Marsabit District, Eastern Province, Kenya, Africa
- Type: Heller, E. 1913 Sep 16. Smithsonian Miscellaneous Collections. 61 (13): 2.
The Lesser Kudu inhabits dry, flat, and densely thicketed areas, as well as woodlands and hilly land. It is rarely observed in open or cleared areas.
(Nowak 1999; Roosevelt and Heller 1914; Walther 1990)
Terrestrial Biomes: savanna or grassland ; scrub forest
Habitat and Ecology
The Lesser Kudu feeds primarily at dusk or at dawn (Roosevelt and Heller 1914) and eats a diverse variety of bush and tree leaves, shoots and twigs, as well as, grasses, herbs, and fruits. It has been reported to be fairly independent of water sources and browses in relatively arid environments.
(Nowak 1999; Roosevelt and Heller 1914; Walther 1990)
Life History and Behavior
Perception Channels: tactile ; chemical
Status: wild: 10.0 years.
Status: captivity: 15.0 years.
Status: wild: 15.0 years.
Lifespan, longevity, and ageing
The Lesser Kudu is a fairly solitary species. Young males stay with their mother for only 1.5-2.0 years, before they leave and travel alone or in small sporadic male groups (Nowak 1999). Young females form small groups with their mother or siblings. The Lesser Kudu becomes sexually mature at 1.25-1.50 years, however, males do not gain social status to reproduce until they reach the age of 4-5 years (Walther 1990). BR>Males perform a shoving match, where they press their heads and horns together and attempt to force their horns down onto the nape of their opponent. Males and females also perform a superiority contest, where the male and female stand fully erect on their hind legs and attempt to push each other over. The larger male usually wins. The males show restraint and are never aggressive towards females, though females have been observed to butt their heads against the males. When the males mount, they lay their neck and head down and onto the females back(Walther 1990). <BR>Each female has its own, independent estrus cycle and is anestrus for only a couple of weeks (Nowak 1999). Gestation ranges from 7.5 to 8.0 months, then the female separates from the group in order to give birth. Only one offspring is produced, weighing 4.0-7.5 kg. 50% of the calves die within the first six months, from disease and predation, and only 25% survive to reach 3 years of age. Males begin horn growth after the first 6-9 months and reach full length after 3 years. The life span of the Lesser Kudu reportedly can reach up to 15 years. <BR
(Harrison and Bates 1991; Nowak 1999; Roosevelt and Heller 1914; Walther 1990)
Range number of offspring: 1 (low) .
Average number of offspring: 1.
Range gestation period: 7.4 to 8.5 months.
Key Reproductive Features: gonochoric/gonochoristic/dioecious (sexes separate); sexual
Average birth mass: 6000 g.
Average number of offspring: 1.
Average age at sexual or reproductive maturity (male)
Sex: male: 504 days.
Average age at sexual or reproductive maturity (female)
Sex: female: 504 days.
Parental Investment: altricial ; post-independence association with parents
Molecular Biology and Genetics
Barcode data: Tragelaphus imberbis
Below is the sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen.
Other sequences that do not yet meet barcode criteria may also be available.
-- end --
Download FASTA File
Statistics of barcoding coverage: Tragelaphus imberbis
Public Records: 1
Specimens with Barcodes: 1
Species With Barcodes: 1
The Lesser Kudu is listed in IUCN as conservation dependent (Nowak 1999). Populations have continued to decline due to hunting or habitat loss caused by human activities or in the case of Tsavo National Park, elephant populations that alter the vegetative landscape.
(Nowak 1999; Walther 1990)
US Federal List: no special status
CITES: no special status
IUCN Red List of Threatened Species: near threatened
IUCN Red List Assessment
Red List Category
Red List Criteria
The Lesser Kudu’s long-term survival prospects would be enhanced by improved protection and management of the relatively few protected areas which support substantial populations. In addition, its value as a trophy animal gives the species high potential for increased revenue generation in the extensive bushlands where it still occurs in good numbers outside national parks and equivalent reserves (East 1999).
Relevance to Humans and Ecosystems
The Lesser Kudu is a hunted game animal, that is used for sport, food, and as a source of money.
Positive Impacts: food ; body parts are source of valuable material
The lesser kudu (Tragelaphus imberbis) is a forest antelope found in East Africa. It is placed in the genus Tragelaphus and family Bovidae. It was first described by the English zoologist Edward Blyth in 1869. The head-and-body length is typically between 110–140 cm (43–55 in). Males reach approximately 95–105 cm (37–41 in) at the shoulder, while females reach 90–100 cm (35–39 in). Males typically weigh 92–108 kg (203–238 lb) and females 56–70 kg (123–154 lb). The females and juveniles have a reddish-brown coat, while the males become yellowish grey or darker after the age of two years. Horns are present only on males. The spiral horns are 50–70 cm (20–28 in) long, and have two to two-and-a-half twists.
A pure browser, the lesser kudu feeds on foliage from bushes and trees, shoots, twigs and herbs. Despite seasonal and local variations, foliage from trees and shrubs constitute 60-80% of the diet throughout the year. The lesser kudu is mainly active at night and during the dawn, and seeks shelter in dense thickets just after the sunrise. The lesser kudu exhibits no territorial behaviour, and fights are rare. While females are gregarious, adult males prefer being solitary. There is no fixed breeding season; births may occur at any time of the year. The lesser kudu inhabits dry, flat and heavily forested regions.
The lesser kudu is native to Ethiopia, Kenya, Somalia, South Sudan, Tanzania and Uganda; while it is extinct in Djibouti. The total population of the lesser kudu has been estimated to be nearly 118,000, with a decreasing trend in populations. One-third of the populations survive in protected areas. Presently, the International Union for Conservation of Nature (IUCN) rates the lesser kudu as "Near Threatened".
Taxonomy and genetics
The scientific name of the lesser kudu is Tragelaphus imberbis. The animal is placed in the genus Tragelaphus and family Bovidae. It was first described by the English zoologist Edward Blyth in 1869. The generic name, Tragelaphus, derives from Greek word Tragos, meaning a male goat, and elaphos, which means a deer; while the specific name imberbis comes from the Latin term meaning unbearded, referring to this kudu's lack of mane. The vernacular name kudu (or koodoo) is the Hottentot name for the greater kudu, a close relative of this species. The term "lesser" denotes the smaller size and lack of mane of this antelope as compared to the greater kudu. In 1912, the genus Ammelaphus was established for just the lesser kudu by American zoologist Edmund Heller, the type species being A. strepsiceros. However, today the lesser kudu is placed in Tragelaphus instead of Ammelaphus.
The lesser kudu has 38 chromosomes, in both males and females. But unlike other tragelaphids, the X chromosome and Y chromosome are compound and fused with autosomes from ancestors having a greater chromosome number.
The lesser kudu is a spiral-horned antelope. The head-and-body length is typically between 110–140 cm (43–55 in). Males reach approximately 95–105 cm (37–41 in) at the shoulder, while females reach 90–100 cm (35–39 in). Males typically weigh 92–108 kg (203–238 lb) and females 56–70 kg (123–154 lb). The bushy tail is 25–40 cm (9.8–15.7 in) long, white underneath and with a black tip at the end.
There are distinct signs of sexual dimorphism in the antelope. The male is considerably larger than the female. The females, as well as juveniles, have a rufous coat, whereas the males become yellowish grey or darker after the age of two years. While the male has a prominent black crest of hair on the neck, this feature is not well-developed in the female. One long white stripe runs along the back, with 11-14 white stripes branching towards the sides. The chest has a central black stripe, and there is no throat beard. A black stripe runs from each eye to the nose and a white one from each eye to the centre of the dark face. A chevron is present between the eyes. The area around the lips is white, the throat has white patches and two white spots appear on each side of the lower jaw. The underparts are completely white, while the slender legs are tawny and have black and white patches. The lesser kudu is characterised by large, rounded ears. Its tracks are similar to the greater kudu's. Females have four teats. The average lifespan is 10 years in the wild, and 15 years in captivity.
Horns are present only on males. The spiral horns are 50–70 cm (20–28 in) long, and have two to two-and-a-half twists. The base circumference is 156–171 cm (61–67 in). The slender horns are dark brown and tipped with white. Male young begin developing horns after six to eight months, which reach full length after three years.
Ecology and behaviour
The lesser kudu is mainly active at night and during the dawn, and seeks shelter in dense thickets just after the sunrise. It can camouflage so well in such dense vegetation that only its ears and tail can indicate its presence. The midday is spent in rest and rumination in shaded areas. The animal spends about 35% of daytime foraging, 36% standing and lying, and 29% in roaming. As the thinnest tragelaphine, the lesser kudu can move through dense vegetation with ease. The lesser kudu is a shy and wary animal. When alarmed, the animal will stand motionless, confirming any danger. If it senses any approaching predator, it will give out a short sharp bark, similar to the bushbuck's. The lesser kudu would then make multiple leaps of up to 2 m (6.6 ft) height with an upraised tail. If captured by the predator, the victim gives a loud bleat.
The lesser kudu is gregarious in nature. There is no distinct leader or any hierarchy in the social structure. There is no territorial behaviour and fights are uncommon. While fighting, the lesser kudu will interlock horns and try pushing one another. Mutual grooming is hardly observed. Unlike most tragelaphines, females can be closely associated for several years. One to three females along with their offspring may form a group. Juvenile males leave their mothers when aged a year-and-a-half, and may form pairs. However, at the age of four to five years, males prefer a solitary lifestyle and avoiding one another, though four to five bulls may share the same home range. Lesser kudu do not usually associate with other animals, except when they feed in the same area.
A pure browser, the lesser kudu feeds on foliage from bushes and trees, shoots, twigs and herbs. It also eats flowers and fruits if available, and takes small proportions of grasses, usually in the wet season. Despite seasonal and local variations, foliage from trees and shrubs constitute 60-80% of the diet throughout the year. Foliage from creepers and vines (such as Thunbergia guerkeana and some species of Cucurbitaceae and Convulvulaceae) form 15-25% of the diet in the wet season. Fruits are consumed mainly in the dry season. Olfactory searching, much in the same posture as grazing, is used to find fallen fruits (like Melia volkensii and Acacia tortilis), while small fruits (like Commiphora species) are directly plucked from trees. The size and structure of its stomach also suggests its primary dependence on browse.
The lesser kudu browses primarily at dusk or at dawn, and is associated with the gerenuk and the impala. The lesser kudu and the gerenuk might compete for evergreen species in the dry season. However, unlike the gerenuk, the lesser kudu rarely prefers Acacia species and does not stand on its hindlegs while feeding. The lesser kudu does not have a great requirement for water, and can browse in arid environments. It eats succulent plants, like the wild sisal, Sansevieria and Euphorbia species in the dry season, and will have a drink when water sources are available.
Both the males and females become sexually mature by the time they are a year-and-a-half old. However, males actually mate after the age of four to five years. Males and females are most reproductive till the age of 14 and 14–18 years respectively, with the maximum age of successful lactation in females being 13–14 years. There is no fixed breeding season; births may occur at any time of the year. A study at Dvůr Králové Zoo (Czech Republic) showed that 55% of the births occurred between September and December. A rutting male will test the urine of any female he encounters, to which the female responds by urinating. Having located a female in estrus, the male follows her closely, trying to rub his cheek on her rump, head, neck and chest. He performs gasping movements with his lips. Finally, the male mounts the female, resting his head and neck on her back, in a similar way as other tragelaphines.
The gestational period is of seven to eight months, after which a single calf is born. A female about to give birth isolates herself from her group, and remains alone for some days afterward. The newborn calf weighs 4–7.5 kg (8.8–16.5 lb). Around 50% of the calves die within the first six months of birth, and only 25% can survive after three years. In a study at Basle Zoo (Switzerland), where 43% of the offspring from captive breeding died before reaching the age of six months, the major causes of high juvenile mortality were found to be the spread of white muscle disease and deficiency of vitamin E and selenium in diets. The herd size, sex, interbreeding and season did not play any role in juvenile mortality. The mother hides her calf while she goes out to feed, and returns mainly in the evening to suckle her young. She checks the calf's identity by sniffing its rump or neck. In the first month, suckling may occur for eight minutes. The mother and calf communicate with low bleats. She licks her offspring, particularly in the perineal region, and may consume its excreta.
Habitat and distribution
The lesser kudu inhabits dry, flat and heavily forested regions. It is closely associated with Acacia and Commiphora thornbush in semi-arid areas of northeastern Africa. The animal avoids open areas and long grass, preferring shaded areas with short grasses instead. Found in woodlands and hilly areas as well, the lesser kudu is generally found at altitudes below 1,200 m (3,900 ft); though they have been recorded at heights of about 1,740 m (5,710 ft) near Mount Kilimanjaro. While individual home ranges of these animals are 0.4–6.7 km2 (4,300,000–72,000,000 sq ft) in size, those of males have an average size of 2.2 km2 (24,000,000 sq ft) and those of females 1.8 km2 (19,000,000 sq ft).
The lesser kudu is native to Ethiopia, Kenya, Somalia, South Sudan, Tanzania and Uganda; while it is extinct in Djibouti. Largely confined to the Horn of Africa today, the species historically ranged from Awash (Ethiopia) southward through southern and eastern Ethiopia, and most parts of Somalia (except the north and the northeast) and Kenya (except the southwest). It also occurred in southeastern Sudan and northeastern and eastern parts of Uganda and Tanzania. The only evidence for its existence in the Arabian peninsula is a single set of horns obtained in 1967 from an individual shot in South Yemen and another in Saudi Arabia.
Threats and conservation
The lesser kudu's shyness and its ability to camouflage itself in dense cover has protected it from the risks of poaching. For instance, the lesser kudu is widespread in the Ogaden region, which is rich in dense bush, despite reckless hunting by local people here. However, rinderpest outbreaks, to which the lesser kudu is highly susceptible, have resulted in a steep decline of 60% in the animal's population in Tsavo National Park in Kenya. Overgrazing, human settlement, loss of habitat are some other threats to the survival of the lesser kudu.
The total population of the lesser kudu has been estimated to be nearly 118,000, with a decreasing trend in populations. The rate of decline has increased to 20% over two decades. Presently, the IUCN rates the lesser kudu as "Near Threatened". Around a third of the population of the lesser kudu occur in protected areas such as Awash, Omo and Mago National Parks (Ethiopia); Bush Bush National Park (Somalia); Tsavo National Park (Kenya); Ruaha National Park and game reserves (Tanzania); though it occurs in larger numbers outside these areas. Population density rarely exceeds 1/km2., and is generally much lower.
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