Panthera leo melanochaita
|Panthera leo melanochaita|
|Male lion in Etosha National Park, Namibia|
|Female lion in Samburu National Reserve, Kenya|
P. l. melanochaita
|Panthera leo melanochaita|
(Ch. H. Smith, 1842)
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 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.
Charles Hamilton Smith described the type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus. 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 bleyenberghi (Lönnberg 1914), a male lion from the Katanga Province of Belgian Congo
- Felis leo roosevelti (Heller 1914), a lion from the Ethiopian Highlands presented to Theodore Roosevelt
- Felis leo nyanzae (Heller 1914), a lion skin from Kampala, Uganda
- 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 Ludwig Zukowsky 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.
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. Based on the results of a genetic analyses, it appears 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 diverged about 50,000 years ago. It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.
Among six samples from captive lions that originated in Ethiopia, five samples clustered with samples from East Africa, but one clustered with samples from the Sahel. For a subsequent phylogeographic study, also eight wild lion samples from the Ethiopian Highlands were included in the analysis using 194 lion sequences from 22 countries. Four wild lion samples from Ethiopia clustered with lion samples from Central Africa, and four with samples from East Africa, indicating that the Great Rift Valley was not a complete barrier to gene flow. Southeastern Ethiopia is therefore considered a genetic admixture zone between Central and East African lions.
Lions 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.
The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. In the 19th and 20th centuries, lion type specimens were described on the basis of mane size and colour. Mane colour varies between tawny, isabelline and light reddish yellow. The Cape lion type specimen had a black mane extending beyond the shoulders and under the belly. Yet, black-maned lions also occur in the Kalahari and eastern Okavango Delta alongside those with a normal tawny colour. Two male lions observed in the border region between Kenya and Tanzania had moderate tufts of hair on the knee joint, and their manes appeared brushed backwards. They were less cobby with longer legs and less curved backs than lions from other African range countries. Male lions from the Ethiopian Highlands had dark and heavy manes with black tips that extended over the whole throat and chest to the forelegs and behind the shoulders. A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well. In ancient Egyptian art, lions are depicted with weak manes. The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations. Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.
In 2002, research in Serengeti National Park revealed that mane darkens with age; its colour and size are influenced by environmental factors like temperature and climate, but also by individual testosterone production, sexual maturity and genetic precondition. Mane length apparently signals fighting success in male–male relationships. 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 have become sexually mature. Males living in the highlands above an altitude of 800 m (2,600 ft) develop heavier manes than lions in the more humid and warmer lowlands of eastern and northern Kenya. The latter have thinner manes, or are even completely maneless.
White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in South Africa. Their whitish fur is a rare morph caused by a double recessive allele. It has normal pigmentation in eyes and skin. White individuals have been occasionally encountered only in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. 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. White lions are selected for breeding in captivity. Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.
Size and weight
Average head-to-body length of male lions is 2.47–2.84 m (97–112 in) with a weight of 148.2–190.9 kg (327–421 lb). Females are smaller and less heavy. The largest known lion measured 3.33 m (10.9 ft). An exceptionally heavy male lion near Mount Kenya weighed 272 kg (600 lb). Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa. In 1936, a man-eating lion shot outside Hectorspruit in Eastern Transvaal weighed about 313 kg (690 lb) and was considered to be the heaviest wild lion. The longest wild lion reportedly was a male shot near Mucusso in southern Angola in 1973.
Distribution and habitat
In East and Southern Africa, lion populations declined in:
- 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 in northeastern Democratic Republic of the Congo.
- 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. As of 2006, 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 3,684 km2 (1,422 sq mi). Lion populations in Kenya and Tanzania are fragmented over 17 patches ranging in size from 86 to 127,515 km2 (33 to 49,234 sq mi).
- 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.
|Range countries||Lion Conservation Units||Area in km2|
|Democratic Republic of Congo||Massif D'itombwe, Luama||8,441|
|Democratic Republic of Congo, Uganda||Queen Elizabeth−Virunga||5,583|
|Uganda||Toro−Semulik, Lake Mburo, Murchison Falls||4,800|
|Kenya||Laikipia−Samburu, Meru and Nairobi National Parks||43,706|
|Kenya, Tanzania||Serengeti−Mara and Tsavo−Mkomazi||75,068|
|Tanzania||Dar-Biharamulo, Ruaha−Rungwa, Mpanga-Kipengere, Tarangire, Wami Mbiki−Saadani, Selous||384,489|
|Mozambique||Cahora Bassa, Gilé, Gorongosa−Marromeu||82,715|
|Mozambique, Zambia||Middle Zambezi||64,672|
|Mozambique, South Africa||Great Limpopo Transfrontier Park||150,347|
|Zambia||Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex||72,569|
|Zambia, Malawi||North−South Luangwa||72,992|
|Botswana, South Africa||Kgalagadi||163,329|
|Angola||Kissama−Mumbondo, Bocoio−Camacuio, Alto Zambeze||393,760|
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.
Behaviour and ecology
In the Serengeti National Park, monitoring of lion prides started in 1966. Between 1966 and 1972, two observed lion prides comprised between seven and 10 females each. Females had litters once in 23 months on average. 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. Male lions in coalitions are closely related. 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.
Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002.
In the 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. 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.
Hunting and diet
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), zebra (Equus burchellii), 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). 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. Faeces of lions collected near waterholes in Hwange National Park also contained remains of climbing mice (Dendromus) and common mice (Mus).
In Botswana's Chobe National Park, lions also prey on 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. In October 2005, lions killed eight elephants aged between one and 11 years, and two of them older than eight years.
Attacks on humans
Several cases of lion attacking people have been documented:
- In the 1890s, two Tsavo Man-Eaters attacked workers during the building of the Uganda Railway. Their skulls and skins are part of the zoological collection of the Field Museum of Natural History in Chicago. The total number of people killed is unclear, but allegedly 135 people fell victim to these lions in less than a year before Colonel Patterson killed them.
- The Njombe lions were a lion pride in Njombe in former Tanganyika, which are thought to have preyed on 1,500 to 2,000 people. They were eventually killed by George Gilman Rushby.
- Between 1990 and 2004, lions killed more than 560 people in Tanzania, mostly during harvest time in crop fields and in areas where natural prey is scarce.
- In February 2018, lions killed a suspected poacher near Kruger National Park.
- In February 2018, Kevin Richardson took three lions for a walk at Dinokeng Game Reserve in South Africa. A lioness pursued an impala for at least 2 km (1.2 mi), and killed a young woman near her car.
- In July 2018, human remains were found in the lion enclosure of a privately owned reserve in South Africa. They were suspected to have been rhino poachers, as they had a high-powered rifle with a silencer, an axe and wire cutters.
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. 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.
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. Private game ranches in South Africa also breed lions for the canned hunting industry.
In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle. 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.
African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes. 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. 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.
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.
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. In addition, about 100 captive lions were registered in ISIS as P. l. krugeri, which derived from lions captured in South Africa. Interest in the Cape lion had led to attempts to conserve possible descendants in Tygerberg Zoo.
Lion populations in Southern and East Africa were referred to by several regional names, including "Katanga lion", "Transvaal lion", "Kalahari lion", "Southeast African lion", and "Southwest African lion", "Masai lion", "Serengeti lion," "Tsavo lion" and "Uganda lion".
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