Panthera leo melanochaita is a lion subspecies in Southern and East Africa. 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. 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. In 2005, a Lion Conservation Strategy was developed for East and southern Africa.
Results of a phylogeographic study indicate that lion populations in southern and eastern Africa form a major clade distinct from lion populations in West Africa, Central Africa and Asia. 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. 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.
The type specimen for P. l. melanochaita was a black-maned lion from the Cape of Good Hope, known as the Cape lion. 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. It has been referred to as the Southern lion, Southern African lion, East-Southern African lion and the "southern subspecies".
Taxonomy
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. In the 19th and 20th centuries, several naturalists described zoological specimens from Southern and East Africa and proposed the following subspecies: Felis leo somaliensis (Noack 1891), based on two lion specimens from Somalia Felis leo massaicus (Neumann 1900), based on two lions killed near Kibaya and the Gurui River in Kenya Felis leo sabakiensis (Lönnberg 1910), based on two male lions from the environs of Mount Kilimanjaro Felis leo roosevelti (Heller 1913), a lion from the Ethiopian Highlands presented to Theodore Roosevelt Felis leo nyanzae (Heller 1913), a lion skin from Kampala, Uganda Felis leo bleyenberghi (Lönnberg 1914), a male lion from the Katanga Province of Belgian Congo Leo leo hollisteri (Joel Asaph Allen 1924), a male lion from the area of Lime Springs, Sotik on the eastern shore of Lake Victoria Leo leo krugeri (Austin Roberts 1929), an adult male lion from the Sabi Sand Game Reserve named in honour of Paul Kruger Leo leo vernayi (Roberts 1948), a male lion from the Kalahari collected by the Vernay-Lang Kalahari Expedition Panthera leo webbensies 1964, two lions from Somalia, one in the Natural History Museum, Vienna that originated in Webi Shabeelle, the other kept in a German zoo that had been imported from the hinterland of Mogadishu.
Dispute over the validity of these purported subspecies continued among naturalists and curators of natural history museums until the early 21st century. In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies. 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.
Pocock subordinated the lion to the genus Panthera in 1930, when he wrote about Asiatic lions. 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. Various authors recognized between seven and 10 African lion subspecies. Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.
In the 1970s, the scientific name P. l. vernayi was considered synonymous with P. l. krugeri. In 1975, Vratislav Mazák hypothesized that the Cape lion evolved geographically isolated from other populations by the Great Escarpment. 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.
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. In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo. Two lion subspecies are now recognised: P. l. melanochaita is understood as comprising lion populations in the contemporary Southern and East African range countries, P. l. leo comprises lion populations in North, West and Central Africa and Asia.
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.
Phylogeny
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. These groups genetically diverged between 50,000 and 200,000 years ago. It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.
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. 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.
Overlap between subspecies
Among six samples of captive lions from Ethiopia, five clustered with samples from East Africa, but one with samples from the Sahel. 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.
Distribution and habitat
In East and Southern Africa, lion populations declined in: In Ethiopia, where lion populations declined since at least the early 20th century due to trophy hunting by Europeans, killing of lions by local people out of fear, for illegal sale of skins and during civil wars. , between seven and 23 lions were estimated to live in Nechisar National Park located in the Ethiopian Highlands. This small protected area is encroached by local people and their livestock. In 2012, lions were documented in cloud forest habitat of Kafa Biosphere Reserve. Somalia since the early 20th century. Intensive poaching since the 1980s and civil unrest posed a threat to lion persistence. Uganda to near extinction in the 20th century. In 2010, the lion population in Uganda was estimated at 408 ± 46 individuals in three protected areas including Queen Elizabeth, Murchison Falls and Kidepo Valley National Parks. Other protected areas in the country probably host less than 10 lions. Lions in Queen Elizabeth National Park form a contiguous population with lions in Virunga National Park. Kenya in the 1990s due to poisoning of lions and poaching of lion prey species. At least 108 lions were killed between 2001 and 2006 in the Amboseli−Tsavo West−Tsavo East National Park network. , there were an estimated 675 lions in the Tsavo national parks, out of the 2,000 total in Kenya. Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of . Lion populations in Kenya and Tanzania are fragmented over 17 patches ranging in size from . Rwanda and Tanzania due to killing of lions during the Rwandan Civil War and ensuing refugee crisis in the 1990s. A small population was present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004. Lions were reintroduced to this national park in 2015. Malawi and Zambia due to illegal hunting of prey species in protected areas. Botswana due to intensive hunting and conversion of natural habitats for settlements since the early 19th century. In Northern Tuli Game Reserve, 19 lions died between 2005 and 2011 due to poaching, trophy hunting and snaring. Namibia due to massive killing of lions by farmers since at least the 1970s. In 2010, the small and isolated lion population in the Kalahari was estimated at 683 to 1,397 individuals in three protected areas, the Kgalagadi Transfrontier Park, the Kalahari Gemsbok and Gemsbok National Parks. South Africa since the early 19th century in the Natal and Cape Provinces south of the Orange River, where the Cape lion population was eradicated by 1860. A few decades later, lions in the Highveld north of the Orange River were also eradicated. In Transvaal, lions occurred historically in the Highveld as well, but were restricted to eastern Transvaal's Bushveld by the 1970s. Between 2000 and 2004, 34 lions were reintroduced to eight protected areas in the Eastern Cape Province, including Addo Elephant National Park. In Venetia Limpopo Nature Reserve, 18 lions were trophy hunted and 11 euthanized between 2005 and 2011.
Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped. Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals. 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 .
North East African clade
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. Range countries Lion Conservation Units Area in km2Ethiopia South Omo, Nechisar, Bale, Welmel-Genale, Awash National Parks, Ogaden 93,274Somalia Arboweerow−Alafuuto 24,527Somalia, Kenya Bushbush−Arawale22,540Kenya Laikipia−Samburu, Meru and Nairobi National Parks 43,706
South East African clade
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.