Panthera leo melanochaita is a
lionsubspecies 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]
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]
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:
Felis leo somaliensis (Noack 1891), based on two lion specimens from
Somalia[12]
Felis leo massaicus (
Neumann 1900), based on two lions killed near
Kibaya and the Gurui River in
Kenya[13]
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
genusPanthera 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]
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.[30]
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.[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
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
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:
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.[40] As of 2009[update], 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.[41] In 2012, lions were documented in
cloud forest habitat of
Kafa Biosphere Reserve.[42]
Somalia since the early 20th century.[43] Intensive
poaching since the 1980s and civil unrest posed a threat to lion persistence.[44][45]
Uganda to near extinction in the 20th century.[46] 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.[47] Lions in
Queen Elizabeth National Park form a contiguous population with lions in
Virunga National Park.[48]
Kenya in the 1990s due to poisoning of lions and poaching of lion prey species.[44] At least 108 lions were killed between 2001 and 2006 in the
Amboseli−
Tsavo West−
Tsavo East National Park network.[49] As of 2006[update], there were an estimated 675 lions in the Tsavo national parks, out of the 2,000 total in Kenya.[49] Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of 3,684 km2 (1,422 sq mi).[50] 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).[51]
Malawi and
Zambia due to illegal hunting of prey species in protected areas.[44]
Botswana due to intensive hunting and conversion of natural habitats for settlements since the early 19th century.[54] In Northern Tuli Game Reserve, 19 lions died between 2005 and 2011 due to poaching, trophy hunting and
snaring.[55]
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[update].[39]
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.[5]
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]
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]
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
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 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
recessiveallele. 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
Tree-climbing lion in Murchison Falls National Park
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
Lionesses hunting a Cape buffalo in the Okavango Delta
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:
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 killed by
George Gilman Rushby.[95]
Between 1990 and 2004, lions killed more than 560 people in Tanzania, mostly during harvest season in crop fields and in areas where natural prey is scarce.[96]
In February 2018, lions killed a suspected poacher near Kruger National Park.[97][98]
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.[99]
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.[100]
Threats
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
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]
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 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]
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