Panthera leo melanochaita

Panthera leo melanochaita
Male lion in Maasai Mara National Reserve, Kenya
Lioness in Etosha National Park, Namibia
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	P. leo
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]

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. ^[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

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:

• 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]
• Felis leo sabakiensis(Lönnberg 1910), based on two male lions from the environs of Mount Kilimanjaro ^[14]
• Felis leo roosevelti(Heller 1913), a lion from the Ethiopian Highlands presented to Theodore Roosevelt ^[15]
• Felis leo nyanzae(Heller 1913), a lion skin from Kampala, Uganda ^[15]
• Felis leo bleyenberghi(Lönnberg 1914), a male lion from the Katanga Province of Belgian Congo ^[16]
• Leo leo hollisteri(Joel Asaph Allen 1924), a male lion from the area of Lime Springs, Sotik on the eastern shore of Lake Victoria ^[17]
• Leo leo krugeri(Austin Roberts 1929), an adult male lion from the Sabi Sand Game Reserve named in honour of Paul Kruger ^[18]
• Leo leo vernayi(Roberts 1948), a male lion from the Kalahari collected by the Vernay-Lang Kalahari Expedition ^[19]
• 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. ^[20]

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]

• 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

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

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]
• Rwanda and Tanzania due to killing of lions during the Rwandan Civil War and ensuing refugee crisis in the 1990s. ^[44] A small population was present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004. ^[52] Lions were reintroduced to this national park in 2015. ^[53]
• 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]
• Namibia due to massive killing of lions by farmers since at least the 1970s. ^[56] 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. ^[57]
• 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. ^[6] A few decades later, lions in the Highveld north of the Orange River were also eradicated. ^[58] In Transvaal, lions occurred historically in the Highveld as well, but were restricted to eastern Transvaal's Bushveld by the 1970s. ^[59] Between 2000 and 2004, 34 lions were reintroduced to eight protected areas in the Eastern Cape Province, including Addo Elephant National Park. ^[60] In Venetia Limpopo Nature Reserve, 18 lions were trophy hunted and 11 euthanized between 2005 and 2011. ^[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

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 countries	Lion Conservation Units	Area in km2
Ethiopia	South Omo, Nechisar, Bale, Welmel-Genale, Awash National Parks, Ogaden	93,274 [61]
Somalia	Arboweerow−Alafuuto	24,527 [4]
Somalia, Kenya	Bushbush−Arawale	22,540 [4]
Kenya	Laikipia−Samburu, Meru and Nairobi National Parks	43,706 [39]

South East African clade

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 countries	Lion Conservation Units	Area 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 Elizabeth−Virunga National Parks	5,583 [48]
Kenya, Tanzania	Serengeti−Mara, Tarangire and Tsavo−Mkomazi	75,068 [63]
Tanzania	Dar-Biharamulo, Ruaha−Rungwa, Mpanga-Kipengere, Wami Mbiki−Saadani, Selous	384,489 [63]
Tanzania, Mozambique	Niassa	177,559 [64]
Mozambique	Cahora Bassa, Gilé, Gorongosa−Marromeu	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	North−South 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

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 countries	Lion Conservation Units	Area in km2
Zimbabwe	Mapungubwe, Bubye	10,033 [39]
Botswana, Zimbabwe	Okavango−Hwange	99,552 [39]
Botswana	Xaixai	12,484 [4]
Angola	Kissama−Mumbondo, Bocoio−Camacuio, Alto Zambeze	393,760 [4]
Angola, Namibia	Etosha−Kunene	123,800 [4]
Namibia	Khaudum−Caprivi	92,372 [4]

Characteristics

Adult male lions in Serengeti National Park

Female lion in the Okavango Delta

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

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

Main article: White lion

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

Tree-climbing lion in Murchison Falls National Park

Lion pride at the Zambezi River

Lions mating in Etosha National Park

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

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:

• 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. ^[93] 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. ^[94]
• 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]

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

Captive lion in Philadelphia Zoo

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

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

• Wild cats in Africa: African leopard  · African golden cat  · Caracal  · Serval  · African wildcat  · Sand cat  · Black-footed cat  · Cheetah

• African Cats
• Born Free
• Elsa the lioness
• The Lion King
• Mapogo lion coalition
• Pride

References

1. Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O'Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z. & Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News. Special Issue 11: 71–73. Archived (PDF) from the original on 30 July 2018. Retrieved 5 August 2019.
2. Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P. & Nowell, K. (2016). "Panthera leo". IUCN Red List of Threatened Species . 2016: e.T15951A115130419.
3. Bauer, H.; Chapron, G.; Nowell, K.; Henschel, P.; Funston, P.; Hunter, L. T.; Macdonald, D. W. & Packer, C. (2015). "Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas". Proceedings of the National Academy of Sciences. 112 (48): 14894–14899. Bibcode:2015PNAS..11214894B. doi: 10.1073/pnas.1500664112 . PMC   4672814 . PMID   26504235.
4. IUCN Cat Specialist Group (2006). Conservation Strategy for the Lion Panthera leo in Eastern and Southern Africa (PDF). Pretoria, South Africa: IUCN. Archived (PDF) from the original on 3 August 2020. Retrieved 1 December 2018.
5. Bertola, L. D.; Jongbloed, H.; Van Der Gaag, K. J.; De Knijff, P.; Yamaguchi, N.; Hooghiemstra, H.; Bauer, H.; Henschel, P.; White, P. A.; Driscoll, C. A. & Tende, T. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports. 6: 30807. Bibcode:2016NatSR...630807B. doi:10.1038/srep30807. PMC   4973251 . PMID   27488946.
6. Mazak, V. (1975). "Notes on the Black-maned Lion of the Cape, Panthera leo melanochaita (Ch. H. Smith, 1842) and a Revised List of the Preserved Specimens". Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen (64): 1–44.
7. Barnett, R.; Sinding, M. H.; Vieira, F. G.; Mendoza, M. L.; Bonnet, M.; Araldi, A.; Kienast, I.; Zambarda, A.; Yamaguchi, N.; Henschel, P. & Gilbert, M. T. (2018). "No longer locally extinct? Tracing the origins of a lion (Panthera leo) living in Gabon". Conservation Genetics. 19 (3): 611–618. doi:10.1007/s10592-017-1039-2. PMC   6448349 . PMID   31007636.
8. Bauer, H.; Chardonnet, P.; Nowell, K. (December 2005), Status and distribution of the lion (Panthera leo) in East and Southern Africa (PDF), Johannesburg, South Africa: East and Southern African lion Conservation Workshop, archived (PDF) from the original on 29 July 2018, retrieved 3 September 2018
9. Hunter, L. & Barrett, P. (2018). "Lion Panthera leo". The Field Guide to Carnivores of the World (2 ed.). London, Oxford, New York, New Delhi, Sydney: Bloomsbury. pp. 46−47. ISBN   978-1-4729-5080-2. Archived from the original on 5 March 2024. Retrieved 27 December 2018.
10. Schofield, A. (2013). White Lion: Back to the Wild. BookBaby. ISBN   978-0-620-57005-3.^{[ permanent dead link ‍]}
11. Smith, C. H. (1842). "Black maned lion Leo melanochaitus". In Jardine, W. (ed.). The Naturalist's Library. Vol. 15 Mammalia. London: Chatto and Windus. p. Plate X, 177.
12. Noack, T. (1891). "Felis leo". Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten. 9 (1): 120.
13. Neumann, O. (1900). "Die von mir in den Jahren 1892–95 in Ost- und Central-Afrika, speciell in den Massai-Ländern und den Ländern am Victoria Nyansa gesammelten und beobachteten Säugethiere". Zoologische Jahrbücher. Abtheilung für Systematik, Geographie und Biologie der Thiere. 13 (VI): 529–562.
14. Lönnberg, E. (1910). "Mammals". In Sjöstedt, Y. (ed.). Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906. Volume 1. Stockholm: P. Palmquists Aktiebolag. pp. 1–58, Plate 1−7.
15. Heller, E. (1913). "New races of carnivores and baboons from equatorial Africa and Abyssinia". Smithsonian Miscellaneous Collections. 61 (19): 1–12.
16. Lönnberg, E. (1914). "New and rare mammals from Congo". Revue de Zoologie Africaine (3): 273–278.
17. Allen, J. A. (1924). "Carnivora Collected By The American Museum Congo Expedition". Bulletin of the American Museum of Natural History. 47: 73–281.
18. Roberts, A. (1929). "New forms of African mammals". Annals of the Transvaal Museum. 21 (13): 82–121.
19. Roberts, A. (1948). "Descriptions of some new subspecies of mammals". Annals of the Transvaal Museum. 21 (1): 63–69.
20. Zukowsky, L. (1964). "Eine neue Löwenrasse als weiterer Beleg für die Verzwergung der Wirbeltierfauna des afrikanischen Osthorns". Milu, Wissenschaftliche und Kulturelle Mitteilungen aus dem Tierpark Berlin (1): 269–273.
21. Allen, G. M. (1939). A Checklist of African Mammals. Vol. 83. Cambridge, Mass. : The Museum. pp. 1–763.{{cite book}}: |work= ignored (help)
22. Lundholm, B. (1952). "A skull of a Cape lioness (Felis leo melanochaitus H. Smith". Annals of the Transvaal Museum (32): 21–24.
23. Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung. 17: 167–280.
24. Wozencraft, W. C. (2005). "Panthera leo". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 546. ISBN   978-0-8018-8221-0. OCLC   62265494.
25. Pocock, R. I. (1930). "The lions of Asia". Journal of the Bombay Natural Historical Society. 34: 638–665.
26. Ellerman, J. R.; Morrison-Scott, T. C. S. (1966). "Subgenus Leo Oken, 1816, (Brehm, 1829)". Checklist of Palaearctic and Indian Mammals 1758 to 1946 (Second ed.). London: British Museum (Natural History). p. 319.
27. Meester, J.; Setzer, H. W. (1977). The mammals of Africa. An identification manual. Washington, DC: Smithsonian Institution Press.
28. Yamaguchi, N. (2000). "The Barbary lion and the Cape lion: their phylogenetic places and conservation" (PDF). African Lion Working Group News. 1: 9–11. Archived from the original (PDF) on 4 March 2016. Retrieved 30 November 2018.
29. Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation" (PDF). Conservation Genetics. 7 (4): 507–514. doi:10.1007/s10592-005-9062-0. S2CID   24190889. Archived from the original (PDF) on 24 August 2006.
30. Manuel, M. d.; Ross, B.; Sandoval-Velasco, M.; Yamaguchi, N.; Vieira, F. G.; Mendoza, M. L. Z.; Liu, S.; Martin, M. D.; Sinding, M.-H. S.; Mak, S. S. T.; Carøe, C.; Liu, S.; Guo, C.; Zheng, J.; Zazula, G.; Baryshnikov, G.; Eizirik, E.; Koepfli, K.-P.; Johnson, W. E.; Antunes, A.; Sicheritz-Ponten, T.; Gopalakrishnan, S.; Larson, G.; Yang, H.; O'Brien, S. J.; Hansen, A. J.; Zhang, G.; Marques-Bonet, T. & Gilbert, M. T. P. (2020). "The evolutionary history of extinct and living lions". Proceedings of the National Academy of Sciences of the United States of America . 117 (20): 10927–10934. doi: 10.1073/pnas.1919423117 . PMC   7245068 . PMID   32366643.
31. Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)" (PDF). Proceedings of the Royal Society B: Biological Sciences. 273 (1598): 2119–2125. doi:10.1098/rspb.2006.3555. PMC   1635511 . PMID   16901830. Archived from the original (PDF) on 8 August 2007.
32. Dubach, J.; Patterson, B. D.; Briggs, M. B.; Venzke, K.; Flamand, J.; Stander, P.; Scheepers, L.; Kays, R. W. (2005). "Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo". Conservation Genetics. 6 (1): 15–24. doi:10.1007/s10592-004-7729-6. S2CID   30414547. Archived from the original on 5 March 2024. Retrieved 29 August 2019.
33. Antunes, A.; Troyer, J. L.; Roelke, M. E.; Pecon-Slattery, J.; Packer, C.; Winterbach, C.; Winterbach, H.; Johnson, W. E. (2008). "The Evolutionary Dynamics of the Lion Panthera leo revealed by Host and Viral Population Genomics". PLOS Genetics. 4 (11): e1000251. doi: 10.1371/journal.pgen.1000251 . PMC   2572142 . PMID   18989457.
34. Dubach, J. M.; Briggs, M. B.; White, P. A.; Ament, B. A.; Patterson, B. D. (2013). "Genetic perspectives on "Lion Conservation Units" in Eastern and Southern Africa". Conservation Genetics. 14 (4): 741–755. doi:10.1007/s10592-013-0453-3. S2CID   751286.
35. O'Brien, S. J.; Martenson, J. S.; Packer, C.; Herbst, L.; de Vos, V.; Joslin, P.; Ott-Joslin, J.; Wildt, D. E. & Bush, M. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions" (PDF). National Geographic Research. 3 (1): 114–124. Archived from the original (PDF) on 2 May 2013.
36. Mazák, J. H. (2010). "Geographical variation and phylogenetics of modern lions based on craniometric data". Journal of Zoology. 281 (3): 194–209. doi:10.1111/j.1469-7998.2010.00694.x.
37. Bertola, L. D.; Van Hooft, W. F.; Vrieling, K.; Uit De Weerd, D. R.; York, D. S.; Bauer, H.; Prins, H. H. T.; Funston, P. J.; Udo De Haes, H. A.; Leirs, H.; Van Haeringen, W. A.; Sogbohossou, E.; Tumenta, P. N.; De Iongh, H. H. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa" (PDF). Journal of Biogeography . 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x. S2CID   82728679. Archived from the original (PDF) on 8 June 2019. Retrieved 30 August 2018.
38. Bertola, L. D.; Tensen, L.; Van Hooft, P.; White, P. A.; Driscoll, C. A.; Henschel, P.; Caragiulo, A.; Dias-Freedman, I.; Sogbohossou, E. A.; Tumenta, P. N.; Jirmo, T. H. (2015). "Autosomal and mtDNA markers affirm the distinctiveness of lions in West and Central Africa". PLOS ONE. 10 (10): e0137975. Bibcode:2015PLoSO..1037975B. doi: 10.1371/journal.pone.0137975 . PMC   4605676 . PMID   26466139.
39. Riggio, J.; Jacobson, A.; Dollar, L.; Bauer, H.; Becker, M.; Dickman, A.; Funston, P.; Groom, R.; Henschel, P.; De Iongh, H.; Lichtenfeld, L. (2013). "The size of savannah Africa: a lion's (Panthera leo) view". Biodiversity and Conservation. 22 (1): 17–35. doi: 10.1007/s10531-012-0381-4 .
40. Yalden, D. W.; Largen, M. J.; Kock, D. (1980). "Catalogue of the Mammals of Ethiopia". Monitore Zoologico Italiano. 13 (1): 169−272. doi: 10.1080/00269786.1980.11758553 .
41. Yirga, G.; Gebresenbet, F.; Deckers, J.; Bauer, H. (2014). "Status of Lion (Panthera leo) and Spotted Hyena (Crocuta crocuta) in Nechisar National Park, Ethiopia". Momona Ethiopian Journal of Science. 6 (2): 127–137. doi: 10.4314/mejs.v6i2.109714 .
42. Gebresenbet, F.; Baraki, B.; Yirga, G.; Sillero-Zubiri, C. & Bauer, H. (2018). "A culture of tolerance: coexisting with large carnivores in the Kafa Highlands, Ethiopia". Oryx. 52 (4): 751–760. doi: 10.1017/S0030605316001356 . Archived from the original on 7 March 2023. Retrieved 20 May 2022.
43. Funaioli, U. and Simonetta, A. M. (1966). "The Mammalian Fauna of the Somali Republic: Status and Conservation Problems". Monitore Zoologico Italiano, Supplemento. 1 (1): 285–347. doi: 10.1080/03749444.1966.10736746 .
44. Chardonnet, P. (2002). "Chapter II: Population Survey" (PDF). Conservation of the African Lion: Contribution to a Status Survey. Paris: International Foundation for the Conservation of Wildlife, France & Conservation Force, USA. pp. 21–101. Archived from the original (PDF) on 10 November 2013.
45. Fagotto, F. (1985). "Larger animals of Somalia in 1984". Environmental Conservation. 12 (3): 260–264. doi:10.1017/S0376892900016015. S2CID   86042501.
46. Treves, A.; Naughton-Treves, L. (1999). "Risk and opportunity for humans coexisting with large carnivores". Journal of Human Evolution. 36 (3): 275–282. doi:10.1006/jhev.1998.0268. PMID   10074384. S2CID   32901765.
47. Omoya, E. O.; Mudumba, T.; Buckland, S. T.; Mulondo, P. & Plumptre, A. J. (2014). "Estimating population sizes of lions Panthera leo and spotted hyaenas Crocuta crocuta in Uganda's savannah parks, using lure count methods" (PDF). Oryx. 48 (3): 394–401. doi: 10.1017/S0030605313000112 . Archived (PDF) from the original on 30 September 2020. Retrieved 30 November 2018.
48. Treves, A.; Plumptre, A. J.; Hunter, L. T. B.; Ziwa, J. (2009). "Identifying a potential lion Panthera leo stronghold in Queen Elizabeth National Park, Uganda, and Parc National des Virunga, Democratic Republic of Congo". Oryx. 43: 60–66. doi: 10.1017/S003060530700124X (inactive 2 November 2024).
49. Frank, L.; Maclennan, S.; Hazzah, L.; Hill, T.; Bonham, R. (2006). Lion Killing in the Amboseli-Tsavo Ecosystem, 2001–2006, and its Implications for Kenya's Lion Population (PDF). Nairobi: Living with Lions. Archived (PDF) from the original on 7 March 2023. Retrieved 30 November 2018.
50. Dolrenry, S.; Hazzah, L.; Frank, L. G. (2016). "Conservation and monitoring of a persecuted African lion population by Maasai warriors". Conservation Biology. 30 (3): 467–475. doi:10.1111/cobi.12703. PMID   27111059. S2CID   3780510.
51. Dolrenry, S.; Stenglein, J.; Hazzah, L.; Lutz, R. S.; Frank, L. (2014). "A metapopulation approach to African lion (Panthera leo) conservation". PLOS ONE. 9 (2): e88081. Bibcode:2014PLoSO...988081D. doi: 10.1371/journal.pone.0088081 . PMC   3914926 . PMID   24505385.
52. Bauer, H.; Van Der Merwe, S. (2004). "Inventory of free-ranging lions Panthera leo in Africa". Oryx. 38 (1): 26–31. doi: 10.1017/S0030605304000055 .
53. Bantlin, D. A. (2017). Movements of Reintroduced Lions in Akagera National Park, Rwanda, in Relation to Prey Abundance (Doctoral dissertation). Madison: University of Wisconsin−Madison.
54. Smithers, R. H. N. (1971). "Panthera leo". The Mammals of Botswana. Pretoria: University of Pretoria. pp. 116–119.
55. Snyman, A.; Jackson, C. R.; Funston, P. J. (2015). "The effect of alternative forms of hunting on the social organization of two small populations of lions Panthera leo in Southern Africa". Oryx. 49 (4): 604–610. doi: 10.1017/S0030605313001336 .
56. Stander, P. E. (1990). "A suggested management strategy for stock-raiding lions in Namibia". South African Journal of Wildlife Research. 20 (2): 37–43.
57. Ferreira, S. M.; Govender, D.; Herbst, M. (2013). "Conservation implications of Kalahari lion population dynamics". African Journal of Ecology. 51 (2): 176–179. doi:10.1111/aje.12003.
58. Guggisberg, C. A. W. (1975). "Lion Panthera leo (Linnaeus, 1758)". Wild Cats of the World. New York: Taplinger Publishing. pp. 138–179. ISBN   978-0-8008-8324-9.
59. Rautenbach, I. L. (1978). "Panthera leo (Linnaeus, 1758)". The Mammals of the Transvaal. Pietermaritzburg: University of Natal. pp. 471–475.
60. Hayward, M. W.; Adendorff, J.; O'Brien, J.; Sholto-Douglas, A.; Bissett, C.; Moolman, L. C.; Bean, P.; Fogarty, A.; Howarth, D.; Slater, R.; Kerley, G. I. (2007). "Practical considerations for the reintroduction of large, terrestrial, mammalian predators based on reintroductions to South Africa's Eastern Cape Province" (PDF). The Open Conservation Biology Journal. 1 (1): 1–11. doi: 10.2174/1874839200701010001 . Archived (PDF) from the original on 9 August 2017. Retrieved 30 November 2018.
61. Riggio, J.; Jacobson, A.; Dollar, L.; Bauer, H.; Becker, M.; Dickman, A.; Funston, P.; Groom, R.; Henschel, P.; de Iongh, H.; Lichtenfeld, L.; Pimm, S. (2013). "The size of savannah Africa: a lion's (Panthera leo) view". Biodiversity Conservation. 22 (1): 17–35. doi: 10.1007/s10531-012-0381-4 .
62. Mudumba, T.; Mulondo, P.; Okot, E.; Nsubuga, M.; Plumptre, A. (2009). National Census of Lions and Hyaenas in Uganda. Strategic Action Plan for Large Carnivore Conservation in Uganda. Kampala: Uganda Wildlife Authority.
63. Mésochina, P.; Mbangwa, O.; Chardonnet, P.; Mosha, R.; Mtui, B.; Drouet, N.; Kissui, B. (2010). Conservation status of the lion (Panthera leo Linnaeus, 1758) in Tanzania. Paris: SCI Foundation, MNRT-WD, TAWISA & IGF Foundation.
64. Chardonnet, P.; Mésochina, P.; Bento, C.; Conjo, D.; Begg, C. l. (2009). Conservation status of the lion (Panthera leo Linnaeus, 1758) in Mozambique (PDF). Maputo, Mozambique: Fondation IGF, Campfire Association, SCI Foundation, The Ratel Trust, Direcção Nacional de Terras e Florestas. Archived (PDF) from the original on 28 April 2020. Retrieved 4 December 2018.
65. Smuts, G. L.; Robinson, G. A.; Whyte, I. J. (1980). "Comparative growth of wild male and female lions (Panthera leo)". Journal of Zoology. 190 (3): 365–373. doi:10.1111/j.1469-7998.1980.tb01433.x.
66. Keet, D. F.; Kriek, N. P. J. & Mills, M. G. L. (1999). The epidemiology of tuberculosis in free-ranging lions in the Kruger National Park (PDF) (Report). South African National Parks. Archived (PDF) from the original on 2 July 2021. Retrieved 13 August 2019.
67. Calder, W. A. (1996). "Skeletal muscle". Size, Function, and Life History. New York: Courier Corporation. pp. 17–21. ISBN   978-0-486-69191-6. Archived from the original on 5 March 2024. Retrieved 13 September 2021.
68. Davis, D. D. (1962). "Allometric relationships in Lions vs. Domestic Cats". Evolution. 16 (4): 505–514. doi: 10.1111/j.1558-5646.1962.tb03240.x .
69. Nowell, K.; Jackson, P. (1996). "African lion" (PDF). Wild Cats: Status Survey and Conservation Action Plan. Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21. ISBN   978-2-8317-0045-8. Archived (PDF) from the original on 7 August 2007. Retrieved 30 November 2018.
70. Wood, G. L. (1982). The Guinness Book of Animal Facts and Feats (Third ed.). Enfield, Middlesex: Guinness Superlatives Ltd. ISBN   978-0-85112-235-9.
71. East African Business Digest. University Press of Africa, with contributions from the Kenya National Chamber of Commerce & Industry. 1963. Archived from the original on 5 March 2024. Retrieved 18 March 2018.
72. Guggisberg, C. A. W. (1963). Simba: The Life of the Lion. Philadelphia, PA: Chilton Books. ASIN   B000OKBJQ0.
73. Borzo, G. (2002). "Unique social system found in famous Tsavo lions". EurekAlert. Archived from the original on 22 February 2014. Retrieved 30 November 2018.
74. Gnoske, T. P.; Celesia, G. G. & Kerbis Peterhans, J. C. (2006). "Dissociation between mane development and sexual maturity in lions (Panthera leo): solution to the Tsavo riddle?". Journal of Zoology. 270 (4): 551–560. doi:10.1111/j.1469-7998.2006.00200.x.
75. West, P. M. & Packer, C. (2002). "Sexual Selection, Temperature, and the Lion's Mane". Science . 297 (5585): 1339–1943. Bibcode:2002Sci...297.1339W. doi:10.1126/science.1073257. PMID   12193785. S2CID   15893512.
76. West, P. M. (2005). "The Lion's Mane: Neither a token of royalty nor a shield for fighting, the mane is a signal of quality to mates and rivals, but one that comes with consequences". American Scientist. 93 (3): 226–235. doi:10.1511/2005.3.226. JSTOR   27858577.
77. Turner, J. A.; Vasicek, C. A.; Somers, M. J. (2015). "Effects of a colour variant on hunting ability: the white lion in South Africa". Open Science Repository Biology: e45011830.
78. McBride, C. (1977). The White Lions of Timbavati. Johannesburg: E. Stanton. ISBN   978-0-949997-32-6.
79. Tucker, L. (2003). Mystery of the White Lions—Children of the Sun God. Mapumalanga: Npenvu Press. ISBN   978-0-620-31409-1.
80. Schaller, G. B. (1972). The Serengeti lion: A study of predator–prey relations. Chicago: University of Chicago Press. ISBN   978-0-226-73639-6.
81. Bertram, B.C.R. (1973). "Lion population regulation". East African Wildlife Journal. 11 (3–4): 215–225. doi:10.1111/j.1365-2028.1973.tb00088.x.
82. Bertram, B. (1975). "Social factors influencing reproduction in wild lions". Journal of Zoology. 177 (4): 463–482. doi:10.1111/j.1469-7998.1975.tb02246.x.
83. Bygott, J. D.; Bertram, B. C. R. & Hanby, J. P. (1979). "Male lions in large coalitions gain reproductive advantages". Nature. 282 (5741): 839–841. Bibcode:1979Natur.282..839B. doi:10.1038/282839a0. S2CID   4249090.
84. Borrego, N.; Ozgul, A.; Slotow, R.; Packer, C. (2018). "Lion population dynamics: do nomadic males matter?". Behavioral Ecology. 29 (3): 660–666. doi: 10.1093/beheco/ary018 .
85. Spong, G. (2002). "Space use in lions, Panthera leo, in the Selous Game Reserve: social and ecological factors". Behavioral Ecology and Sociobiology. 52 (4): 303–307. doi:10.1007/s00265-002-0515-x. S2CID   37631550.
86. Elliot, N. B.; Cushman, S. A.; Macdonald, D. W. & Loveridge, A. J. (2014). "The devil is in the dispersers: predictions of landscape connectivity change with demography". Journal of Applied Ecology. 51 (5): 1169–1178. doi: 10.1111/1365-2664.12282 .
87. Valeix, M.; Loveridge, A. J.; Davidson, Z.; Madzikanda, H.; Fritz, H. & Macdonald, D. W. (2010). "How key habitat features influence large terrestrial carnivore movements: waterholes and African lions in a semi-arid savanna of north-western Zimbabwe". Landscape Ecology. 25 (3): 337–351. doi:10.1007/s10980-009-9425-x. S2CID   24083943.
88. Patterson, B. D., Kasiki, S. M., Selempo, E. and Kays, R. W. (2004). "Livestock predation by lions (Panthera leo) and other carnivores on ranches neighboring Tsavo National Parks, Kenya". Biological Conservation. 119 (4): 507–516. CiteSeerX   10.1.1.884.8117 . doi:10.1016/j.biocon.2004.01.013.
89. Hayward, M. W.; Kerley, G. I. H. (2005). "Prey preferences of the lion (Panthera leo)" (PDF). Journal of Zoology. 267 (3): 309–322. CiteSeerX   10.1.1.611.8271 . doi:10.1017/S0952836905007508. Archived (PDF) from the original on 27 June 2015. Retrieved 30 November 2018.
90. Davidson, Z., Valeix, M., Van Kesteren, F., Loveridge, A.J., Hunt, J.E., Murindagomo, F. and Macdonald, D.W. (2013). "Seasonal diet and prey preference of the African lion in a waterhole-driven semi-arid savanna". PLOS ONE. 8 (2): e55182. Bibcode:2013PLoSO...855182D. doi: 10.1371/journal.pone.0055182.g001 . PMC   3566210 . PMID   23405121.
91. Joubert, D. (2006). "Hunting behaviour of lions (Panthera leo) on elephants (Loxodonta africana) in the Chobe National Park, Botswana". African Journal of Ecology. 44 (2): 279–281. doi:10.1111/j.1365-2028.2006.00626.x.
92. Power, R. J.; Compion, R. X. S. (2009). "Lion predation on elephants in the Savuti, Chobe National Park, Botswana". African Zoology. 44 (1): 36–44. doi:10.3377/004.044.0104. S2CID   86371484. Archived from the original on 31 August 2018. Retrieved 30 November 2018.
93. Kerbis Peterhans, J. C. and Gnoske, T. P. (2001). "The Science of 'Man-Eating*' Among Lions Panthera leo With a Reconstruction of the Natural History of the 'Man-Eaters of Tsavo'". Journal of East African Natural History. 90 (1): 1–40. doi: 10.2982/0012-8317(2001)90[1:TSOMAL]2.0.CO;2 .
94. Patterson, J. H. (1907). The man-eaters of Tsavo and other East African adventures. London: Macmillan and Co.
95. Rushby, G. G. (1965). No More the Tusker. London: W. H. Allen. Archived from the original on 5 March 2024. Retrieved 10 September 2020.
96. Packer, C.; Ikanda, D.; Kissui, B.; Kushnir, H. (2005). "Lion attacks on humans in Tanzania" (PDF). Nature. 436 (7052): 927–928. Bibcode:2005Natur.436..791S. doi:10.1038/436791a. PMID   16094358. S2CID   4312804. Archived from the original (PDF) on 1 July 2020. Retrieved 6 December 2018.
97. "South African lions eat 'poacher', leaving just his head". The BBC. 2018. Archived from the original on 23 February 2018. Retrieved 25 February 2018.
98. Haden, A. (2018). "Suspected poacher mauled to death by lions close to Kruger National Park". The South African. Archived from the original on 24 February 2018. Retrieved 25 February 2018.
99. Torchia, C. (2018). "Lion kills woman at refuge of South African 'lion whisperer'". The Washington Post . Archived from the original on 15 January 2019. Retrieved 28 February 2018.
100. "Lions eat 'rhino poachers' on South African game reserve". BBC . 2018. Archived from the original on 17 July 2018. Retrieved 19 July 2018.
101. Packer, C.; Brink, H.; Kissui, B. M.; Maliti, H.; Kushnir, H.; Caro, T. (2011). "Effects of trophy hunting on lion and leopard populations in Tanzania". Conservation Biology. 25 (1): 142–153. doi:10.1111/j.1523-1739.2010.01576.x. PMID   20825444. S2CID   205658449.
102. Williams, V. L.; Loveridge, A. J.; Newton, D. J.; Macdonald, D. W. (2017). "A roaring trade? The legal trade in Panthera leo bones from Africa to East-Southeast Asia". PLOS ONE. 12 (10): e0185996. Bibcode:2017PLoSO..1285996W. doi: 10.1371/journal.pone.0185996 . PMC   5655489 . PMID   29065143.
103. Schroeder, R. A. (2018). "Moving Targets: The 'Canned'Hunting of Captive-Bred Lions in South Africa". African Studies Review. 61 (1): 8–32. doi:10.1017/asr.2017.94. S2CID   150275315.
104. "'Herdsmen' poison lions, vultures — but not in Nigeria". The Cable. 2018. Archived from the original on 25 February 2018. Retrieved 24 February 2018.
105. Kamoga, J. (2018). "East African lions dying of poisoning". The Observer . Archived from the original on 22 February 2018. Retrieved 24 February 2018.
106. Winter, S. (2018). "Lion MASSACRE as six big cats die after eating 'poison'". Daily Express . Archived from the original on 24 February 2018. Retrieved 24 February 2018.
107. "Zimbabwe's 'iconic' lion Cecil killed by hunter". BBC News. 2015. Archived from the original on 28 July 2015. Retrieved 3 August 2015.
108. "Xanda, son of Cecil the lion, killed by hunter in Zimbabwe". BBC News. 2017. Archived from the original on 21 July 2017. Retrieved 20 July 2017.
109. Midlane, N. (2013). The conservation status and dynamics of a protected African lion Panthera leo population in Kafue National Park, Zambia. Cape Town: University of Cape Town.
110. Cushman, S. A.; Elliot, N. B.; Bauer, D.; Kesch, K.; Bahaa-el-din, L.; Bothwell, H.; et al. (2018). "Prioritizing core areas, corridors and conflict hotspots for Lion conservation in Southern Africa". PLOS ONE. 13 (7): e0196213. Bibcode:2018PLoSO..1396213C. doi: 10.1371/journal.pone.0196213 . PMC   6033387 . PMID   29975694.
111. Bruche, S.; Gusset, M.; Lippold, S.; Barnett, R.; Eulenberger, K.; Junhold, J.; Driscoll, C. A.; Hofreiter, M. (2012). "A genetically distinct lion (Panthera leo) population from Ethiopia". European Journal of Wildlife Research. 59 (2): 215–225. doi:10.1007/s10344-012-0668-5. S2CID   508478.
112. Tefera, M. (2003). "Phenotypic and reproductive characteristics of lions (Panthera leo) at Addis Ababa Zoo". Biodiversity & Conservation. 12 (8): 1629–1639. doi:10.1023/A:1023641629538. S2CID   24543070.
113. Hogarth, C.; Butler, N. (2004). "Animal Symbolism (Africa)". In Walter, M. N. (ed.). Shamanism: An Encyclopedia of World Beliefs, Practices, and Culture, Volume 1 . Bloomsbury Academic. pp. 3–6. ISBN   978-1-57607-645-3.
114. Lynch, P. A. (2004). "Lion". African Mythology A to Z. Infobase Publishing. p.  63. ISBN   978-0-8160-4892-2.
115. Brakefield, T. (1993). Big Cats: Kingdom of Might . Voyageur Press. p.  44. ISBN   978-0-89658-329-0.
116. Jackson, D. (2010). "Introduction". Lion. London: Reaktion Books. pp. 1–21. ISBN   978-1-86189-735-0. Archived from the original on 5 March 2024. Retrieved 30 November 2018.

External links

• Carrington, D. (2018). "Lion poaching: the brutal new threat to Africa's prides". The Guardian.
• "Lion (Panthera leo ssp. melanochaita)". ECOS Environmental Conservation Online System. 2016.
• $0.5m funding to stop the decline in the population of African lions
• What Will It Take to Save the East African Lion from Extinction? Hunting or Herding?
• Lions in East Africa
• Recovering population of Zimbabwean African lions show low genetic diversity
• PetaPixel, September 2018: Lioness Steals Photographer's Canon DSLR and Gives It to Her Cubs
• National Geographic Wildlife Watch, June 2018: As Tigers Become Rarer, Poachers Are Targeting Lions
• Lobengula the South African guardian lion
• What Happened to Angola's 1,000 Lions? Archived 2018-11-24 at the Wayback Machine
• Holding the line for lions in Mozambique
• Giant Lions Once Prowled East Africa, 200,000-Year-Old Skull Reveals
• BBC Earth: Lions take down an adult elephant
• The Savuti Lions of the Chobe National Park
• A Zambian lion stirs Archived 2018-08-07 at the Wayback Machine
• Shamba the South African lion
• Known for escapes, South African lion becomes a father