Panthera leo melanochaita

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Panthera leo melanochaita
Die pure Kraft - Löwe im Etosha-Nationalpark.JPG
Male lion in Etosha National Park, Namibia
Lioness Samburu 2.jpg
Female lion in Samburu National Reserve, Kenya
Scientific classification e
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
(Ch. H. Smith, 1842)
Synonyms[1]

formerly:

  • P. l. bleyenberghi
  • P. l. krugeri
  • P. l. vernayi
  • P. l. massaica
  • P. l. nyanzae

Panthera leo melanochaita is a lion subspecies in Southern and East Africa.[2] 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.[3] 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.[4] In 2005, a Lion Conservation Strategy was developed for East and Southern 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]

Taxonomic history[edit]

Lions shot in Kenya's Sotik Plains in 1909

Charles Hamilton Smith described the type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus.[7] 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][17][18][19][1] In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies.[14][17] 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.[17]

Pocock subordinated the lion to the genus Panthera in 1930, when he wrote about Asiatic lions.[20] 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.[21] Various authors recognized between seven and 10 African lion subspecies.[19] Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.[22]

In the 1970s, the scientific name P. l. vernayi was considered synonymous with P. l. krugeri.[19] 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.[23][24]

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.[1] In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo.[3] Two lion subspecies are now recognised:[2]

  • 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.

Genetic research[edit]

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. 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.[25] It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.[24][26][27][28][29][30][31]

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.[28] 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.[31]

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.[32]

Characteristics[edit]

Adult male lions in Serengeti National Park, Tanzania
Male lion in Narok County, southern Kenya
A male lion in Amboseli National Park, Kenya
White lion

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft.[33] 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.[34] The Cape lion type specimen had a black mane extending beyond the shoulders and under the belly.[7] Yet, black-maned lions also occur in the Kalahari and eastern Okavango Delta alongside those with a normal tawny colour.[35] 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.[9] 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.[12] A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well.[11] In ancient Egyptian art, lions are depicted with weak manes.[36] The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.[37] Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.[19][6]

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.[38] 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.[39]

White lion[edit]

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.[40] White lions are selected for breeding in captivity.[41] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[42]

Size and weight[edit]

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).[33] An exceptionally heavy male lion near Mount Kenya weighed 272 kg (600 lb).[43] Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa.[44] 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.[45]

Distribution and habitat[edit]

The Serengeti and Maasai Mara National Parks are a lion stronghold in East Africa with a stable lion population[46]

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.[5] Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals.[48][63]

Range countries Lion Conservation Units Area in km2
Democratic Republic of Congo Massif D'itombwe, Luama 8,441[5]
Democratic Republic of Congo, Uganda Queen ElizabethVirunga 5,583[52]
Uganda Toro−Semulik, Lake Mburo, Murchison Falls 4,800[64]
Somalia Arboweerow−Alafuuto 24,527[5]
Somalia, Kenya BushbushArawale 22,540[5]
Kenya LaikipiaSamburu, Meru and Nairobi National Parks 43,706[63]
Kenya, Tanzania SerengetiMara and TsavoMkomazi 75,068[46]
Tanzania Dar-Biharamulo, RuahaRungwa, Mpanga-Kipengere, Tarangire, Wami Mbiki−Saadani, Selous 384,489[46]
Tanzania, Mozambique Niassa 177,559[65]
Mozambique Cahora Bassa, Gilé, GorongosaMarromeu 82,715[65]
Mozambique, Zambia Middle Zambezi 64,672[65]
Mozambique, South Africa Great Limpopo Transfrontier Park 150,347[65]
Zambia Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex 72,569[63]
Zambia, Malawi NorthSouth Luangwa 72,992[63]
Malawi Kasungu, Nkhotakota 4,187[63]
Zimbabwe Mapungubwe, Bubye 10,033[63]
Botswana, Zimbabwe OkavangoHwange 99,552[63]
Botswana Xaixai 12,484[5]
Botswana, South Africa Kgalagadi 163,329[63]
Angola Kissama−Mumbondo, BocoioCamacuio, Alto Zambeze 393,760[5]
Angola, Namibia EtoshaKunene 123,800[5]
Namibia KhaudumCaprivi 92,372[5]

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.[63]

Behaviour and ecology[edit]

Lions mating in Etosha National Park, Namibia
Lionesses hunting a Cape buffalo in the Okavango Delta, Botswana
Male lion and cub feeding on a Cape buffalo, Sabi Sand Game Reserve, South Africa

In the Serengeti National Park, monitoring of lion prides started in 1966.[66] Between 1966 and 1972, two observed lion prides comprised between seven and 10 females each. Females had litters once in 23 months on average.[67] 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.[68] Male lions in coalitions are closely related.[69] 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.[70]

Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002.[71]

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.[72] 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.[73]

Hunting and diet[edit]

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).[74] 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.[66] Faeces of lions collected near waterholes in Hwange National Park also contained remains of climbing mice (Dendromus) and common mice (Mus).[75]

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.[76] In October 2005, lions killed eight elephants aged between one and 11 years, and two of them older than eight years.[77]

Attacks on humans[edit]

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.[78] 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.[79]
  • 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.[80]
  • 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.[81]
  • In February 2018, lions killed a suspected poacher near Kruger National Park.[82][83]
  • 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.[84]
  • 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.[85]

Threats[edit]

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.[3] 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.[53]

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.[86] Private game ranches in South Africa also breed lions for the canned hunting industry.[87]

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle.[88] 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.[89][90]

In 2015 and 2017, two male lions, Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park.[91][92]

In Zambia's Kafue National Park, uncontrolled bushfires and hunting of lions and prey species makes it difficult for the lion population to recover. Cub mortality in particular is high.[93]

Conservation[edit]

African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes.[3] 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.[5] 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.[3]

In captivity[edit]

Captive lion in Philadelphia Zoo

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.[37][94]

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.[24] Interest in the Cape lion had led to attempts to conserve possible descendants in Tygerberg Zoo.[95][96]

Regional names[edit]

Lion populations in Southern and East Africa were referred to by several regional names, including "Katanga lion", "Transvaal lion", "Kalahari lion",[11][14][15] "Southeast African lion", and "Southwest African lion",[97] "Masai lion", "Serengeti lion,"[66] "Tsavo lion"[98] and "Uganda lion".[19]

See also[edit]

References[edit]

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