Panthera leo melanochaita

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Panthera leo melanochaita
Male Lion on Rock.jpg
Male lion in Maasai Mara National Reserve, Kenya
Lioness on the prowl.jpg
Lioness in Etosha National Park, Namibia
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
Subspecies:
P. l. melanochaita
Trinomial name
Panthera leo melanochaita
(Smith, 1842)
Synonyms [1]
  • P. l. bleyenberghi
  • P. l. krugeri
  • P. l. hollisteri
  • P. l. massaica
  • P. l. nyanzae

Panthera leo melanochaita is a lion subspecies in Southern and East Africa. [1] In this part of Africa, lion populations are regionally extinct in Lesotho, Djibouti and Eritrea, and are threatened by loss of habitat and prey base, killing by local people in retaliation for loss of livestock, and in several countries also by trophy hunting. [2] Since the turn of the 21st century, lion populations in intensively managed protected areas in Botswana, Namibia, South Africa and Zimbabwe have increased, but declined in East African range countries. [3] In 2005, a Lion Conservation Strategy was developed for East and Southern Africa. [4]

Contents

Results of a phylogeographic study indicate that lion populations in southern and eastern Africa are forming a major clade distinct from lion populations in West Africa, Central Africa and Asia. [5] In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group subsumed lion populations according to the major clades into two subspecies, namely P. l. leo and P. l. melanochaita. [1] Within P. l. melanochaita three subclades are clearly distinguishable. One from northeastern Africa, another one from southwestern Africa and a third one from southeastern Africa. [5]

The type specimen for P. l. melanochaita was a black-maned lion from the Cape of Good Hope, known as the Cape lion. [6] Phylogeographic analysis of lion samples from Gabon and the Republic of the Congo indicate their close genetic relation to P. l. melanochaita samples from Namibia and Botswana. [7] It has been referred to as the Southern lion, Southern African lion, East-Southern African lion [8] and the "southern subspecies". [9] [10]

Taxonomy

The Big Game of Africa (1910) - Lioness Sotik Plains 1909.png
The Big Game of Africa (1910) - Male Lion Sotik Plains 1909.png
Lions shot in Kenya's Sotik Plains in 1909

Felis (Leo) melanochaitus was the scientific name proposed by Charles Hamilton Smith in 1842 who described a lion specimen from South Africa's Cape Province. [11] In the 19th and 20th centuries, several naturalists described zoological specimens from Southern and East Africa and proposed the following subspecies:

Dispute over the validity of these purported subspecies continued among naturalists and curators of natural history museums until the early 21st century. [6] [21] [22] [23] [24] In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies. [18] [21] In 1939, the American zoologist Allen also recognized F. l. bleyenberghi, F. l. krugeri and F. l. vernayi as valid subspecies in Southern Africa, and F. l. hollisteri, F. l. nyanzae and F. l. massaica as valid subspecies in East Africa. [21]

Pocock subordinated the lion to the genus Panthera in 1930, when he wrote about Asiatic lions. [25] Ellerman and Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African P. l. leo and the Asiatic P. l. persica. [26] Various authors recognized between seven and 10 African lion subspecies. [23] Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa. [27]

In the 1970s, the scientific name P. l. vernayi was considered synonymous with P. l. krugeri. [23] In 1975, Vratislav Mazák hypothesized that the Cape lion evolved geographically isolated from other populations by the Great Escarpment. [6] In the early 21st century, Mazák's hypothesis about a geographically isolated evolution of the Cape lion was challenged. Genetic exchanges between populations in the Cape, Kalahari and Transvaal Province regions and farther east are considered having been possible through a corridor between the Great Escarpment and the Indian ocean. [28] [29]

In 2005, the authors of Mammal Species of the World recognized P. l. bleyenberghi, P. l. krugeri, P. l. vernayi, P. l. massaica, P. l. hollisteri and P. l. nyanzae as valid taxa. [24] In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo. [2] Two lion subspecies are now recognised: [1]

Genome-wide data of a wild-born historical lion sample from Sudan clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. l. melanochaita. This result indicates that the taxonomic position of lions in Central Africa may require revision. [30]

Phylogeny

Range map including proposed clades and the two subspecies (P. l. leo and P. l. melanochaita) according to genetic research Lion subspecies distribution3.png
Range map including proposed clades and the two subspecies (P. l. leo and P. l. melanochaita) according to genetic research

Since the beginning of the 21st century, several phylogenetic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Scientists analysed between 32 and 480 lion samples from up to 22 countries. Results of genetic analyses indicate that the species comprises two main evolutionary groups, one in Southern and East Africa, and the other in the northern and eastern parts of its historical range. [31] [32] [33] [34] These groups genetically diverged between 50,000 and 200,000 years ago. [35] [5] It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups. [29] [33] [36] [37] [38] [5]

Lion samples from Gabon's Batéké Plateau National Park and Odzala-Kokoua National Park in Republic of the Congo were found to be genetically closely related to lion samples from Namibia and Botswana. [7] A phylogenetic analysis of lion samples from Africa and Asia showed that they shared a common ancestor probably between 98,000 and 52,000 years ago. Samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This indicates that Central Africa was a melting pot of lion populations after they had become isolated. They possibly migrated through corridors in the Nile Basin during the early Holocene. [30]

Overlap between subspecies

Lion from Ethiopia at Bronx Zoo Annual report - New York Zoological Society (1914) (18433145331).jpg
Lion from Ethiopia at Bronx Zoo

Among six samples of captive lions from Ethiopia, five clustered with samples from East Africa, but one with samples from the Sahel. [37] For a subsequent phylogeographic study, eight wild lion samples from the Ethiopian Highlands were included in the DNA sequencing analysis using 194 lion samples from 22 countries. Four of these samples clustered with samples from Central Africa, and four with samples from East Africa, indicating that the Great Rift Valley, Ethiopia was not a complete barrier to gene flow. Southeastern Ethiopia is therefore considered a genetic admixture zone between Central and East African lions. [5]

Distribution and habitat

Lion cubs Serengeti.jpg
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Lionesses with cubs, Selous Game Reserve (28) (28436353633).jpg
The Serengeti and Maasai Mara National Parks and Selous Game Reserve are lion strongholds in East Africa with stable lion populations [39]

In East and Southern Africa, lion populations declined in:

Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped. [4] Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals. [44] [39] The LCUs Ruaha−Rungwa, Serengeti−Mara, Tsavo−Mkomazi and Selous in East Africa, as well as Luangwa, Kgalagadi, Okavango−Hwange, Mid−Zambezi, Niassa and Greater Limpopo in Southern Africa are currently considered lion strongholds. These LCUs host more than 500 individuals each, and the population trend is stable there as of 2012. [39]

North East African clade

A juvenile male from the northeastern clade at Samburu Lion male with scanty mane at Samburu NR 2.jpg
A juvenile male from the northeastern clade at Samburu

Lions, which can be grouped into the northeastern clade are found in Somalia, Northern Kenya and Ethiopia, with a larger hybridization zone to the Northern lion subspecies (P. l. leo) in Ethiopia. [5]

Range countriesLion Conservation UnitsArea in km2
EthiopiaSouth Omo, Nechisar, Bale, Welmel-Genale, Awash National Parks, Ogaden 93,274 [61]
Somalia Arboweerow−Alafuuto 24,527 [4]
Somalia, Kenya BushbushArawale 22,540 [4]
Kenya LaikipiaSamburu, Meru and Nairobi National Parks 43,706 [39]

South East African clade

A lion male from the southeastern clade in the Serengeti Lion in Serengeti protected area.jpg
A lion male from the southeastern clade in the Serengeti

Lions, which can be grouped into the southeastern clade are found in Southern Kenya, Western DRC, Tanzania, Mozambique, Malawi, Zambia, southern Namibia and South Africa, with a larger hybridization zone to the southwestern lion clade in the Kruger National Park area (Great Limpopo Transfrontier Park). The classification of lions in Uganda is not clarified, since there were no samples included in the most comprehensive recent study. [5]

Range countriesLion Conservation UnitsArea in km2
Uganda Murchison Falls, Toro−Semulik, Lake Mburo 4,800 [62]
Democratic Republic of Congo Itombwe Nature Reserve, Luama 8,441 [4]
Democratic Republic of Congo, Uganda Queen ElizabethVirunga National Parks 5,583 [48]
Kenya, Tanzania SerengetiMara, Tarangire and TsavoMkomazi 75,068 [63]
TanzaniaDar-Biharamulo, RuahaRungwa, Mpanga-Kipengere, Wami Mbiki−Saadani, Selous 384,489 [63]
Tanzania, Mozambique Niassa 177,559 [64]
Mozambique Cahora Bassa, Gilé, GorongosaMarromeu 82,715 [64]
Mozambique, Zambia Middle Zambezi 64,672 [64]
Mozambique, South Africa Great Limpopo Transfrontier Park 150,347 [64]
Zambia Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex 72,569 [39]
Zambia, Malawi NorthSouth Luangwa 72,992 [39]
Malawi Kasungu, Nkhotakota 4,187 [39]
Botswana, South Africa Kgalagadi 163,329 [39]

South West African clade

Lion in Etosha National Park, Namibia Die pure Kraft - Lowe im Etosha-Nationalpark.JPG
Lion in Etosha National Park, Namibia

Lions, which can be grouped into the southwestern clade are found in Angola, Northern Namibia, Northern Botswana and western Zimbabwe, extending southeast into the Tuli block. a There is a larger hybridization zone to the southeastern clade around the Kruger National Park. [5]

Range countriesLion Conservation UnitsArea in km2
Zimbabwe Mapungubwe, Bubye 10,033 [39]
Botswana, Zimbabwe OkavangoHwange 99,552 [39]
Botswana Xaixai 12,484 [4]
Angola Kissama−Mumbondo, BocoioCamacuio, Alto Zambeze 393,760 [4]
Angola, Namibia EtoshaKunene 123,800 [4]
Namibia KhaudumCaprivi 92,372 [4]

Characteristics

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Adult male lions in Serengeti National Park
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Female lion in the Okavango Delta
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Lion cubs in Zimbabwe

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. [58]

Size and weight

Average head-to-body length of male lions is 2.47–2.84 m (8 ft 1 in – 9 ft 4 in) with a weight ranging from 150–225 kg (331–496 lb) averaging 187.5 kg (413 lb) in Southern Africa and 145.4–204.7 kg (321–451 lb) averaging 174.9 kg (386 lb) in East Africa. Females average 83–165 kg (183–364 lb) in Southern Africa and 90–167.8 kg (198–370 lb) in East Africa. [65] Males in northern Kruger National Park weighed 200.01 kg (440.9 lb) on average, whereas females weighed 143.52 kg (316.4 lb) on average, and males in southern Kruger National Park weighed 186.55 kg (411.3 lb) on average and females weighed 118.37 kg (261.0 lb), though there was an outbreak of tuberculosis in southern park at the time. [66] Skeletal muscles make up 58.8% of the lion's body weight. [67] [68]

The largest known lion measured 3.35 m (11.0 ft) in length and weighed 375 kg (825 lb). [58] An exceptionally heavy male lion near Mount Kenya weighed 272 kg (600 lb). [69] The longest wild lion reportedly was a male shot near Mucusso National Park in southern Angola in 1973. In 1936, a man-eating lion shot in 1936 in eastern Transvaal weighed about 313 kg (690 lb), and was considered to have been one of the heaviest wild lions. [70] In 1963, two lions in Tanzania weighed 320 and 360 kg (700 and 800 lb) after killing several livestock. [71]

Manes

A maneless lion in Tsavo East National Park, Kenya Maneless lion from Tsavo East National Park.png
A maneless lion in Tsavo East National Park, Kenya

In the 19th and 20th centuries, lion type specimens were described on the basis of their mane size and colour. Mane colour varies from sandy, tawny, isabelline, light reddish yellow to dark brown and black. [11] [15] [16] [72] [54] Mane length varies from short to extending to knee joints and under the belly. [23] Lions without a mane were observed in the Tsavo area. [73]

Mane development is related to age: older males have more extensive manes than younger ones; manes continue to grow up to the age of four to five years, long after lions become sexually mature. Males living in the Kenyan highlands above elevations of 800 m (2,600 ft) develop heavier manes than lions in the more humid and warmer lowlands of eastern and northern Kenya. [74] Average ambient temperature, nutrition and testosterone influence the colour and size of the mane. Its length is an indicator for age and fighting ability of the lion. In Serengeti National Park, female lions favour males with dense and dark manes as mates. [75] [76]

White lion

A cub in Kruger National Park Kruger Cub.jpg
A cub in Kruger National Park

White lions have occasionally been encountered in and around South Africa's Kruger National Park and the adjacent Timbavati Private Game Reserve. Their whitish fur is a rare morph caused by a double recessive allele. It has normal pigmentation in eyes and skin. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five different prides between 2007 and 2015. [77] White lions were selected for breeding in captivity. [78] They have been bred in South African camps for use as trophies to be killed during canned hunts. [79]

Behaviour and ecology

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Tree-climbing lion in Murchison Falls National Park
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Lion pride at the Zambezi River
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Lions mating in Etosha National Park
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A lioness with her cub in Botswana

In Serengeti National Park, monitoring of lion prides started in 1966. [80] Between 1966 and 1972, two observed lion prides comprised between seven and 10 females each. Females had litters once in 23 months on average. [81] Litters contained two to three cubs. Of 87 cubs born until 1970, only 12 reached the age of two years. Cubs died due to starvation in months when large prey was not available, or following take-over of the prides by new males. [82] Male lions in coalitions are closely related. [83] Between 1974 and 2012, 471 coalitions comprising 796 male lions entered a study area of 2,000 km2 (770 sq mi). Of these, 35 coalitions included male lions that were born in the area but had left and returned after about two years of absence. Nomadic coalitions became resident at between 3.5 and 7.3 years of age. [84]

The lion population of Selous Game Reserve has been surveyed since 1996. Lion prides avoided acacia woodlands and preferred habitats near water courses with short grasses, where also prey species gathered. Two or more prides shared home ranges. [85]

In Kavango–Zambezi Transfrontier Conservation Area, lions have been monitored since 1999. In 2003, 50 lions were radio-collared in Hwange National Park and tracked until 2012. Results show that adult male and female lions preferred grassland and shrubland habitat, but avoided woodlands and areas with high human density. By contrast, subadult dispersing male lions avoided grasslands and shrublands, but moved in human-dominated areas to a larger extent. Hence, dispersing lions are more vulnerable to coming into conflict with humans than adult lions. [86] In the semi-arid savanna of Zimbabwe's Hwange National Park, 19 lions were radio-collared and tracked between 2002 and 2007. Both female and male lions moved foremost within 2 km (1.2 mi) of waterholes in all seasons. [87]

Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002. [88]

Hunting and diet

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Lionesses hunting a Cape buffalo in the Okavango Delta
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Lions feeding on a giraffe in Kruger National Park

Lions usually hunt in groups and prey foremost on ungulates such as gemsbok (Oryx gazella), Cape buffalo (Syncerus caffer), blue wildebeest (Connochaetes taurinus), giraffe (Giraffa camelopardalis), common eland (Tragelaphus oryx), greater kudu (T. strepsiceros), nyala (T. angasii), roan antelope (Hippotragus equinus), sable antelope (H. niger), plains zebra (Equus quagga), bushpig (Potamochoerus larvatus), common warthog (Phacochoerus africanus), hartebeest (Alcephalus buselaphus), common tsessebe (Damaliscus lunatus), waterbuck (Kobus ellipsiprymnus), kob (K. kob) and Thomson's gazelle (Eudorcas thomsonii). Their prey is usually in the range of 190–550 kg (420–1,210 pounds). [89] In the Serengeti National Park, lions were observed to also scavenge on carrion of animals that were killed by other predators, or died from natural causes. They kept a constant lookout for circling vultures, apparently being aware that vultures indicate a dead animal. [80] Faeces of lions collected near waterholes in Hwange National Park also contained remains of climbing mice ( Dendromus ) and common mice ( Mus ). [90]

In Botswana's Chobe National Park, lions also prey on young and subadult African bush elephants (Loxodonta africana). They successfully attacked 74 elephants between 1993 and 1996, of which 26 were older than nine years, and one bull over 15 years old. [91] In October 2005, a pride of up to 30 lions killed eight African bush elephants that were between four and eleven years old. [92]

Attacks on humans

Several cases of lion attacking people have been documented:

Threats

Taxidermy Lion Trophy Seized by U.S. Fish and Wildlife Service Miami (37402001470).jpg
Taxidermied lion trophy seized by U.S. Fish and Wildlife Service in 2017

In Africa, lions are threatened by pre-emptive killing or in retaliation for preying on livestock. Prey base depletion, loss and conversion of habitat have led to a number of subpopulations becoming small and isolated. Trophy hunting has contributed to population declines in Botswana, Namibia, Zimbabwe and Zambia. [2] It is the primary cause for a decline of lion populations in Tanzania's Selous Game Reserve and Katavi National Park. [101] Although lions and their prey are officially protected in Tsavo National Parks, they are regularly killed by local people, with over 100 known lion killings between 2001 and 2006. [49]

Between 2008 and 2013, bones and body parts from at least 2621 individual lions were exported from South Africa to Southeast Asia, and another 3437 lion skeletons between 2014 and 2016. Lion bones are used to replace tiger bones in traditional Asian medicines. [102] Private game ranches in South Africa also breed lions for the canned hunting industry. [103]

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle. [104] In February 2018, the carcasses of two male and four female lions were found dead in Ruaha National Park, and were suspected to have died of poisoning. [105] [106]

In 2015 and 2017, two male lions, Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park. [107] [108]

Uncontrolled bushfires and hunting of lions and prey species in Zambia's Kafue National Park make it difficult for the lion population to recover. Cub mortality in particular is high. [109]

Conservation

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Lion cubs in Botswana

African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes. [2] IUCN regional offices and many wildlife conservation organisations cooperated to develop a Lion Conservation Strategy for Eastern and Southern Africa in 2006. The strategy envisages to maintain sufficient habitat, ensure a sufficient wild prey base, make lion-human coexistence sustainable and reduce factors that lead to further fragmentation of populations. [4] Local communities in several Southern African lion range countries generate significant income through wildlife tourism, which is a strong incentive for their support of conservation measures. [2]

Establishing corridors between protected areas is important for facilitating dispersal of lions. Makgadikgadi Pans National Park and Central Kalahari Game Reserve are key dispersal areas in Southern Africa. [110]

In captivity

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Captive lion in Philadelphia Zoo
Tourist petting tame lions, Livingstone, Zambia LionWalk.jpg
Tourist petting tame lions, Livingstone, Zambia

At the beginning of the 21st century, the Addis Ababa Zoo kept 16 adult lions. It is assumed that their ancestors, five males and two females, were caught in southwestern Ethiopia as part of a zoological collection for Emperor Haile Selassie I. [111] [112]

In 2006, eight captive lions were registered under the name P. l. massaicus, and 23 as P. l. nubicus from Tanzania by the International Species Information System; about 100 captive lions were registered as P. l. krugeri, which derived from lions captured in South Africa. [29]

In 2012, samples of lions kept at Sana'a Zoo in Yemen were found to cluster with those of lions from East and Southern Africa. [111]

Cultural significance

The Coat of arms of South Africa featured a lion from 1910 to 1932 Coat of arms of the South African Republic.svg
The Coat of arms of South Africa featured a lion from 1910 to 1932

The lion is an animal symbol in shamanistic rituals of the Nuer people. In other East African cultures, it symbolizes laziness. [113] Scars inflicted by lions are regarded as a sign of courage among the Masai people. [114] The name 'Simba' is a Swahili word for the lion, which also means 'aggressive', 'king' and 'strong'. [115]

Regional names

Lion populations in Southern and East Africa were referred to by several regional names, including Katanga lion, Transvaal lion, Kalahari lion, [16] [18] [19] Southeast African lion, and Southwest African lion, [116] Masai lion, Serengeti lion, [80] Tsavo lion [73] and Uganda lion. [23] It has also been referred to as 'Eastern-Southern African lion', [31] [8] 'Southern lion', [33] [30] and as 'southern subspecies'. [9]

See also

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The African leopard is the nominate subspecies of the leopard, native to many countries in Africa. It is widely distributed in most of sub-Saharan Africa, but the historical range has been fragmented in the course of habitat conversion. Leopards have also been recorded in North Africa as well.

<span class="mw-page-title-main">Arabian leopard</span> Subspecies of mammals

The Arabian leopard is the smallest leopard subspecies. It was described in 1830 and is native to the Arabian Peninsula, where it was widely distributed in rugged hilly and montane terrain until the late 1970s. Today, the population is severely fragmented and thought to decline continuously. Previously in 2008, an estimated 45–200 individuals in three isolated subpopulations were restricted to western Saudi Arabia, Oman and Yemen. However, as of 2023, it is estimated that 100–120 in total remains, with 70-84 mature individuals, in Oman and Yemen, and it is possibly extinct in Saudi Arabia. The current population trend is suspected to be decreasing.

Tsavo West National Park is located in the Coast Province of Kenya. The park covers an area of 9,065 square kilometres. The A109 road Nairobi-Mombasa and a railway divides it from the adjoining Tsavo East National Park. Together with adjoining ranches and protected areas, they comprise the Tsavo Conservation Area. Tsavo West is a more popular destination on account of its magnificent scenery, Mzima Springs, rich and varied wildlife, good road system, rhino reserve, rock climbing potential and guided walks along the Tsavo River. The park is operated by Kenya Wildlife Service.

<span class="mw-page-title-main">African bush elephant</span> Species of mammal

The African bush elephant, also known as the African savanna elephant, is one of two extant African elephant species and one of three extant elephant species. It is the largest living terrestrial animal, with bulls reaching a shoulder height of up to 3.96 m and a body mass of up to 10.4 t.

<span class="mw-page-title-main">East African cheetah</span> Subspecies of carnivore

The East African cheetah, is a cheetah population in East Africa. It lives in grasslands and savannas of Tanzania, Kenya, Uganda and Somalia. The cheetah inhabits mainly the Serengeti ecosystem, including Maasai Mara, and the Tsavo landscape.

<span class="mw-page-title-main">Addis Ababa Zoo</span> Zoological park in Addis Ababa, Ethiopia

Addis Ababa Zoo is a zoological park in Addis Ababa, Ethiopia.

<span class="mw-page-title-main">Maneless lion</span> Male lions with a gene polymorphism

The term "maneless lion" or "scanty mane lion" often refers to a male lion without a mane, or with a weak one.

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