User:Punetor i Rregullt5/sandbox/African lion

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African lion
Male
Masai Mara, Kenya
Southern African lioness and cub near Otjiwarongo, Namibia
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
Subspecies:
P. l. leo and melanochaita
Trinomial name
Panthera leo leo and melanochaita
(Linnaeus and CH. H. Smith 1758, 1842)
Roughly, the distribution of lions in Africa

African lions are a genetically diverse population of lions in Africa which are recognised as being divided into two overlapping subspecies as of 2017:[1][2]

The status of lions in the

homogeneous,[1] being divided into different clades.[4]

Formerly described subspecies

Northwest Africa
Subspecies Description
Barbary lion (P. l. leo), also called the "Atlas lion", "Berber lion" or "North African lion" Formerly found in the
nominate lion subspecies from North Africa. It is extinct in the wild due to excessive hunting; the last, known Barbary lion in the wilderness was killed in Morocco in 1920.[5][6] This was regarded as being one of the largest subspecies,[7] with reported lengths of 3.0–3.3 m (9.8–10.8 ft) and weights of more than 200 kg (440 lb) for males. Besides West and certain Central African lions, it is more closely related to the Asiatic lion than to other African lions.[4] A number of animals in captivity are likely to be Barbary lions,[8] particularly the 90 animals descended from the Moroccan Royal collection at Rabat Zoo.[9]

North Africa: Algeria, Libya, Morocco and Tunisia

West Africa
Subspecies Description
Senegal lion (P. l. senegalensis), also known as "
West African lion
"
Found in
critically endangered as of 2015.[12]

West Africa: Benin, Burkina Faso, Ghana,[13] Mali, Nigeria, Niger and Senegal

Gambian lion (P. l. gambianus) Formerly found in the
Gambia.[14]
Central Africa
Subspecies Description
Cameroon lion
(P. l. kamptzi)
Found in Cameroon and the region south of Lake Chad, in Central or Western Africa.[15]
Northeast Congo lion (P. l. azandica), or simply the "
Congo lion
"
Found in northeastern parts of the Congo, adjacent to Uganda.[10]

Central Africa: Democratic Republic of the Congo[16]

Northeast or East Africa
Subspecies Description
Nubian lion (P. l. nubica) From Nubia in Northeast Africa.[17]
Somali lion (P. l. somaliensis syn. P. l. webbiensis From Somaliland or Somalia, East Africa.[18][19]
Masai lion
(P. l. massaica)
Found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique;[11] a local population is known as the "Tsavo lion".

East Africa: Kenya, Tanzania, and Uganda

Ethiopian lion (P. l. roosevelti syn. P. l. abyssinica), also known as "Abyssinian lion" and "Addis Ababa lion" 15 captive lions in the
Haile Selassie I of Ethiopia.[21]

Northeast Africa: Ethiopia

Kilimanjaro lion (P. l. sabakiensis) From the northern vicinity of Mount Kilimanjaro in East Africa.[22]
Ugandan lion (P. l. nyanzae) Found in Uganda, East Africa.[15]
Sotik lion (P. l. hollisteri), also known as "Hollister's lion" or "Lake Victoria lion" Found on the eastern bank of Lake Victoria in Kenya, East Africa.[15]
Southern Africa
Subspecies Description
Cape lion (P. l. melanochaita) Formerly found from the
Natal, South Africa.[23]
Katanga lion (P. l. bleyenberghi), also known as the "Angola lion", "Bleyenbergh's lion" or "Southwest African lion" Found in southwestern Africa. It is among the largest populations of African lions. The type specimen was from Katanga in what in what used to be the Belgian Congo in central Africa.[18][24]

Central Africa: Formerly

Congo-Kinshasa
)

Southern Africa: Angola, Botswana, Namibia, Zambia, and Zimbabwe[11]

Kruger lion
(P. l. krugeri), also known as the "Southeast African lion", "South African lion" or "Transvaal lion"
Found in the Transvaal region of southeast Africa, including Kruger National Park.[11]

Southern Africa: (

Swaziland, Zimbabwe
)

Kalahari lion (P. l. vernayi) Found in the
Kalahari Region of Southern Africa.[15]

Northern subspecies

New York Zoological Gardens
, 1897

The Northern lion

nominate subspecies of the lion, which is present in the northern portion of Africa (particularly in West and north of Central Africa).[2][27] It is regionally extinct in North Africa. In West and Central Africa, lions are restricted to fragmented and isolated populations, most of them declining.[28][29] The West African population is listed as Critically Endangered on the IUCN Red List; this population is isolated and comprises fewer than 250 mature individuals.[30] The Asiatic lion is considered to belong to this subspecies, due to its relationship with North, West and Central African lions.[2]

Results of a

IUCN Cat Specialist Group subsumed lion populations to two subspecies, namely P. l. leo and P. l. melanochaita.[2]

Taxonomic history

A lion from

Linnaeus in 1758.[31] Due to the location of Algeria in Africa, the term "Northern lion" was coined for lions living in the northern portion of the continent in 1865.[25][26] In the 19th and 20th centuries, several lion specimens
from Africa were described and proposed as subspecies:

In the following decades, there has been much debate among zoologists on the validity of proposed subspecies:

  • In 1939, Glover Morrill Allen recognized Felis leo kamptzi as valid taxa among ten lion subspecies.[23][17]
  • Three decades later, John Ellerman and Terence Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African (P. l. leo) and Asiatic lions (P. l. persica).[36]
  • Some authors considered P. l. nubicus a valid subspecies and synonymous with P. l. massaica.[17][18][37]
  • Some authors considered P. l. azandicus synonymous with P. l. massaicus and P. l. somaliensis, and P. l. kamptzi synonymous with P. l. senegalensis.[17][18]
  • In 2005, Wallace Christopher Wozencraft recognized P. l. kamptzi, P. l. bleyenberghi and P. l. azandica as valid taxa.[38]
  • In 2016, IUCN Red List assessors used P. l. leo for all African populations.[39]
  • In 2017, lion populations in North, West and Central Africa and Asia were subsumed to P. l. leo by the Cat Classification Task Force of the IUCN Cat Specialist Group, based on results of genetic research on lion samples.[2]

Genetic research

Rough 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

species comprises two evolutionary groups, one in the northern and eastern parts of its historical range, and the other in Southern and East Africa that diverged between 245,000 and 50,000 years ago. They assume that tropical rainforest and the East African Rift constituted major barriers between the two groups.[40][41][42][43][4]
Based on this assessment, lion taxonomy has been revised as comprising two subspecies:[2]

  • P. l. leo in the northern and eastern regions of its historical and contemporary distribution;
  • P. l. melanochaita in the contemporary southern and East African range countries.

In a comprehensive study about the evolution of lions, 357 samples of 11 lion populations were examined, including some

mtDNA grouping (lineage III) that also included Asiatic samples. This scenario was well in line with theories on lion evolution: lineage III developed in East Africa and traveled north and west in the first wave of lion expansions about 118,000 years ago. It apparently broke up into haplotypes H5 and H6 within Africa, and then into H7 and H8 in West Asia.[41]

Western and Central clades

Results of

phylogenetic data of 194 lion samples from 22 different countries revealed that Central and West African lions form a phylogeographic group that probably diverged about 186,000–128,000 years ago from the melanochaita group in East and Southern Africa.[4]

The West African population is distributed south of the

Gambia, Guinea-Bissau, Mali, Sierra Leone, Ivory Coast, and Togo, and possibly extinct in Guinea.[30]
The largest West African subpopulation of between 246 and 466 individuals survives in the WAP-Complex, a large system of protected areas formed mainly by W, Arli, and Pendjari National Parks in Burkina Faso, Benin, and Niger.[44][45]

The range of the Central African lion clade reaches from the lower

Siniaka-Minia Faunal Reserve and Zakouma and Aouk National Parks, but have been extirpated in Manda National Park
. Lions may still be present in pastoral rangelands and mountain ranges outside protected areas.[28] In 2004, the lion population in the country was estimated at maximum 225 individuals.[27] The following table shows estimates of lion population sizes between 2002 and 2012:

Range countries Area used in km2 Estimated no. of individuals
Bénoué National Park complex, Cameroon 14,682 200[48]
Waza National Park in Cameroon 1,452 17[49]
Central African Republic 339,418 about 1,297[47]
Garamba National Park and Domaine Chasse Bili Uere, Democratic Republic of the Congo 115,671 175−320[29][28]
Southeastern Chad 133,408 400[29]
Southwestern Sudan 331,834 375[29]
Gambela
National Parks, South Sudan and Ethiopia
106,941 500[29]
Omo National Park, Sudan and Ethiopia 22,483 200[29]
other protected and non-protected areas, Ethiopia 70,759 96[29]
Total 1,136,648 3,260−3,305

Contemporary lion distribution and habitat quality in

savannahs of West and Central Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[50] Educated guesses for size of populations in these LCUs ranged from 3,274 to 3,909 individuals between 2002 and 2012.[28][29]

Range countries Lion Conservation Units Area in km2
Senegal, Mali, Guinea-Bissau, Guinea Niokolo-Koba National Park 90,384[29]
Guinea National Park of Upper Niger 613[29]
Benin, Burkina Faso, Niger W-Arly-Pendjari Complex 29,403[29]
Benin three unprotected areas 6,833[29]
Nigeria
Yankari National Park and Kainji National Park
6,551[29]
Cameroon Waza and Bénoué National Parks 16,134[49]
Central African Republic eastern part of the country; Bozoum and Nana Barya Faunal Reserves 339,481[47]
Chad southeastern part 133,408[29]
Democratic Republic of Congo
Bili Uere
115,671[50]
South Sudan, Sudan 331,834[50]
South Sudan, Ethiopia Boma-Gambella 106,941
Ethiopia South 93,274[29]

Southern subspecies

scientific name Panthera leo hollisteri.[38][19]

The Southern lion (Panthera leo melanochaita),

regionally extinct in Lesotho, Djibouti and Eritrea.[39] 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.[53] They 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.[39]

The

type specimen for P. l. melanochaita was a black-maned lion from the Cape of Good Hope, known as the Cape lion, and consequently, the scientific name was initially meant for it. The lion population in this part of South Africa is extinct.[54]

Taxonomic history

type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus.[55] It was referred to as the "Southern relative" of the North African lion.[25][26] In the 19th and 20th centuries, several naturalists described specimens from Southern and East Africa and proposed subspecies
, including:

Dispute over the validity of these purported subspecies continued among naturalists and curators of natural history museums until the early 21st century.[54][38][23][62][17]

In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies.[59][23][62][63] 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.[23]

classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.[66]

In the 1970s, the

In 2005, the authors of

P. l. leo.[39] In 2017, the Cat Classification Task Force of the Cat Specialist Group reduced the number of valid lion subspecies in Southern and Southeast Africa to one, namely P. l. melanochaita.[2]

Genetic research

type specimen for the southern subspecies (P. l. melanochaita). Genetic research has shown it to be closely related to extant Southern African lions.[67]

Since the beginning of the 21st century, several

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.[68] It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.[3][41][69][43][4] Based on this assessment, the species comprises two recognised subspecies:[2]

The two groups were in contact in

West and certain Central African lions. In 9 of 19 lion samples from Ethiopia, haplotypes of the Central African lion group were found, indicating that the Great Rift Valley was not a complete barrier to gene flow; southeastern Ethiopia is considered a genetic admixture zone between Central and East African lions.[4]

Since 2005, several

Transvaal region in South Africa.[70]
Results of phylogeographic studies support the notion of lions in Southern Africa being genetically close, but distinct from populations in West and North Africa and Asia.[41][43] Based on the analysis of samples from 357 lions from 10 countries, it is thought that lions migrated from Southern Africa to East Africa during the Pleistocene and Holocene eras.[41]

A phenotypic and DNA analysis was conducted using samples from 15 captive lions in the Addis Ababa Zoo and from six wild lion populations. Results showed that the captive lions were genetically similar to wild lions from Cameroon and Chad, but with little signs of inbreeding.[21]

Lions samples from Gabon's Batéké Plateau National Park and Odzala-Kokoua National Park in Republic of the Congo were found to be related to the Southern lion clade.[3]

Northeastern clade

Kidepo Valley National Park in northern Uganda, where a lion is hiding in the grass. Lions in Uganda are connected to lions in Central Africa, and thus may be mixed between the northern and southern subspecies.[4]

The range of the Northeastern clade outside the admixture zone is confined to Somalia and northern and central Kenya.[4] Already in the 1980s, the lion population in Somalia had greatly declined due to poaching and was restricted to woodlands in the southern part of the country.[71] In northern Kenya, lions had been observed near Kavirondo, near Lake Manyara and in the Tanga Region in the late 19th century.[57] By the 21st century, lion populations in northern Kenya have been fragmented.[72]

Range countries Area used in km2 Estimated no. of individuals
Laikipia-Samburu complex in Kenya 35,511 271[29]
Meru in Kenya 7,365 40[29]
Arawale complex in Kenya and Somalia 22,540 750[29]
Arboweerow-Alafuuto in Somalia 24,527 175[29]
Total 2,254,067,403 km2 1,236

Southern / Eastern clade

This is the clade with the largest remaining populations. The range of this clade extends from southern Kenya, southern Uganda and the Virunga area in the Democratic Republic of the Congo southward to the Cape of Good Hope, excluding only the western parts of Southern Africa.[4]

The following complexes are considered lion strongholds of the Southern/Eastern clade:[29]

Lions in Queen Elizabeth National Park, which form a contiguous population with lions in Virunga National Park in the northeastern part of the Democratic Republic of the Congo,[29][73][27] do belong to the Southern Eastern clade.[4] 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.[74] As of 2006, there were an estimated 675 lions in the Tsavo area, out of the 2,000 total in Kenya.[75] Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of 3,684 km2 (1,422 sq mi).[72]

A small population is present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004.[27]

The population in South Africa's former

Eastern Cape Province, including Addo Elephant National Park
.

Range countries Area used in km2 Estimated no. of individuals
Virunga and Queen Elizabeth National Park in CAR and Uganda 5,583 210[29]
Lake Mburo in Uganda 373 3[29]
Luama Hunting Reserve in DRC 5,197 <50[29]
Itombwe Massif
in DRC
3,244 <50[29]
North West Tansania 4,703 105[29]
Ruaha-Rungwa in Tanzania 195,993 3,779[29]
Mpanga Kipengere in Tanzania 958 14[29]
Swaga Swaga in Tanzania 7,242 102[29]
Serengeti-Mara in Tanzania and Kenya 35,852 3,673[29]
Nairobi in Kenya 830 <30[29]
Tsavo-Mkomazi in Kenya and Tanzania 39,216 880[29]
Tarangire In Tanzania 28,771 731[29]
Wami Mbiki-Saadani in Tanzania 8,787 136[29]
Selous in Tanzania 138,035 7,644[29]
Niassa in Mozambique, Tanzania 177,559 1,573[29]
Liuwa Plains in Zambia 3,866 4[29]
Kafue in Zambia 58,898 386[29]
Nsumbu in Zambia 5,650 <50[29]
Luangwa National Park in Zambia 72,992 574[29]
Kasungu in Malawi 2,341 4[29]
Nkhotakota in Malawi 1,846 18[29]
Kgalagadi in South Africa and Botswana 163,329 800[29]
Mid-Zambezi in Mozambique, Zambia, Zimbabwe and Mozambique 64,672 755[29]
Tete South of Cahora Bassa, Gile and Gorongosa-Marromeu in Mozambique 13,612, 22,322, 46,781 59, 45, 229[29]

Limpopo admixture zone

Male at Kruger National Park, South Africa

The area of the Kruger National Park, which is part of the Great Limpopo Transfrontier Park, is an admixture zone between the Southern-Eastern and the Southwestern clade. This area is a lion stronghold with about 2,300 lions.[29]

Southwestern clade

Lioness in Etosha National Park, Namibia

The only stronghold of the Southwestern clade is in the western parts of the Kavango–Zambezi Transfrontier Conservation Area, including Okavango Delta and Hwange National Park[77][29] Another important reserve for this clade is the Etosha National Park.[29] Lions are considered regionally extinct in the southwestern part of the Democratic Republic of the Congo.[39][5]

In Gabon, the presence of lions in Batéké Plateau National Park was doubtful in 2010.

phylogenetic analysis and compared with tissue samples of lions from Gabon and Republic of the Congo that were killed in the 20th century. Results indicate that this individual is closely related to the ancestral lion population of the area, and that its DNA shows a typical Southern lion haplotype. It is considered possible that this lion dispersed to the area from Namibia or Botswana.[3]

In the Republic of the Congo, the Odzala-Kokoua National Park was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area, so that now, the species is considered

locally extinct in the country.[80]

Range countries Area used in km2 Estimated no. of individuals
Southwestern clade
Kissama-Mumbondo in Angola 4,593 <10[29]
Bocoio-Camucuio in Angola 22,005 55[29]
SE Angola 386,962 1,905[29]
Sioma Ngwezi in Zambia 4,155 <50[29]
Etosha-Kunene in Namibia 123,800 455[29]
Khaudum-Caprivi in Namibia 92,372 150[29]
Xaixai in Botswana 12,484 75[29]
Okavango-Hwange 99,552 2,300[29]
Greater Mapungubwe in Botswana, South Africa and Zimbabwe 5,158 25[29]
Bubye in Zimbabwe 4,875 200[29]
Total XXX XXX

Northeast African population

Male at Addis Ababa Zoo, Ethiopia

In

trinomina such as Panthera leo nubica, Panthera leo roosevelti and Panthera leo somaliensis, and names such as "Abyssinian lion", "Egyptian lion", "Nubian lion", "Somali lion" and "Sudan lion",[81] depending on the locality of occurrence.[18][15][82]

Genetic analyses and taxonomic history

Young male Nubian lion in New York Zoo, 1903[83]
Abyssinian lion in New York Zoo, 1914[84]

In the

scientific names of the North[23][38] and East African lions.[18][82]

A test done in 2012 on 15 lions at

captive lions were genetically different to wild lions in other parts of East Africa, but similar to wild lions from Cameroon and Chad.[21][20]

Among six samples from captive lions which were of Ethiopian origin, five samples clustered with other East African samples, but one clustered with Sahelian samples.[43] For a subsequent study, also eight additional samples from wild lions from the Ethiopian Highlands were included in a subsequent analysis. Three of them clustered with the Central African lion clade, and five with other East African samples. The Ethiopian Highlands east and west of the Rift Valley was therefore assumed by scientists to have been a zone of genetic admixture between the two phylogeographic groups.[4]

Results of a

National Museum of Natural History (France) that originated in the Nubian part of Sudan. In terms of mitochondrial DNA, it grouped with lion skull samples from the Central African Republic, Ethiopia and the northern part of the Democratic Republic of the Congo.[3]

A phylogeographic analysis of

In 2016, analysis of the genomes of lions showed that there had been a basal split between lions in northern and southern parts of Africa. Lions in northern Central Africa belong to the northern clade, and those in Southeast Africa belonged to the southern clade. Lions samples from Ethiopia were shown to be related to both the South-East African and Central African groups, indicating an overlap between these groups there.[4]

In 2017, the Cat Classification Taskforce of the

Cat Specialist Group subsumed lion populations in West, Central and North Africa to P. l. leo, and those in East Africa to P. l. melanochaita. At the same time, it was stated that these two subspecies overlap in Ethiopia.[2]

One of the largest lion populations in Ethiopia is found in

Alatash National Park close to the international border with Sudan.[86][87][88]

Other parts of Ethiopia, which still have lions fall into the admixture zone. These are Omo and Bale Mountains National Parks, the ara around the Chew Bahir and Turkana lakes, and the Webi Shabeelle area.[27] In 2009, a small group of less than 23 lions were estimated in Nechisar National Park located in the Great Rift Valley. This small protected area in the Ethiopian Highlands is encroached by local people and their livestock.[89]

Lions of northern Uganda have not been analysed genetically,[4] and might belong to the Northern subspecies. In northern Uganda, lions are present in Kidepo Valley and Murchison Falls National Parks.[27][29]

Central African population

Central clade of the northern subspecies[4] (P. l. leo).[2]

The Central African lion

subpopulation, and is fragmented into small and isolated groups since the 1950s.[28][27]

Distribution and habitat

The Serengeti and Maasai Mara National Parks are a lion stronghold in East Africa with a stable lion population[47]

In Northeast Africa, lions may inhabit a variety of habitats, excepting dense

banks of the Blue and White Niles, and far away into desert areas.[81]

Currently, not much is known about the distribution and sizes of populations in

The southern lion was originally found from Ethiopia and Uganda in the north to the Cape of Good Hope in the south. Supported by genetic research, the border between the Southern and Northern subspecies runs through Ethiopia. Southeastern Ethiopia is considered a genetic admixture zone between the two groups. Within the Southern lion, genetic research identified three clades. These are the Northeastern, East-Southern and Southwestern subclade.[4]

In East and Southern Africa, the population of lions declined in:

  • Somalia since the early 20th century.[92]
  • Uganda to near extinction in the 20th century.[93]
  • Kenya in the 1990s due to poisoning of lions and poaching of lion prey species.[28]
  • Rwanda and Tanzania due to killing of lions during the Rwandan Civil War and ensuing refugee crisis in the 1990s.[28]
  • Malawi and Zambia due to illegal hunting of prey species in protected areas.[28]
  • Botswana due to intensive hunting and conversion of natural habitats for settlements since the early 19th century.[94]
  • Namibia due to massive killing of lions by farmers since at least the 1970s.[95]
  • Natal and Cape Provinces south of the Orange River, where the Cape lion population was eradicated by 1860.[54] A few decades later, lions in the Highveld north of the Orange River were also eradicated.[96] In the Transvaal, lions occurred historically in the Highveld as well, but were restricted to eastern Transvaal's Bushveld by the 1970s.[97]

Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[51] Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals.[28][29]

Range countries Lion Conservation Units Area in km2
Democratic Republic of Congo
Massif D'itombwe, Luama
8,441[51]
Democratic Republic of Congo, Uganda Queen Elizabeth-Virunga 5,583[98]
Uganda 4,800[99]
Somalia Arboweerow-Alafuuto 24,527[51]
Somalia, Kenya Bushbush-Arawale 22,540[51]
Kenya Laikipia-Samburu, Meru and Nairobi National Parks 43,706[29]
Kenya, Tanzania
Mara and Tsavo-Mkomazi
75,068[47]
Tanzania Dar- 384,489[47]
Tanzania, Mozambique Niassa 177,559[100]
Mozambique Cahora Bassa, Gilé, Gorongosa-Marromeu 82,715[100]
Mozambique, Zambia Middle Zambezi 64,672[100]
Mozambique, South Africa Great Limpopo Transfrontier Park 150,347[100]
Zambia Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex 72,569[29]
Zambia, Malawi North-South Luangwa 72,992[29]
Malawi Kasungu, Nkhotakota 4,187[29]
Zimbabwe Mapungubwe, Bubye 10,033[29]
Botswana, Zimbabwe Okavango-Hwange 99,552[29]
Botswana Xaixai 12,484[51]
Botswana, South Africa Kgalagadi 163,329[29]
Angola Kissama-Mumbondo, Bocoio-Camacuio, Alto Zambeze 393,760[51]
Angola, Namibia Etosha-Kunene 123,800[51]
Namibia Khaudum-Caprivi 92,372[51]

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 as lion strongholds. These LCUs host more than 500 individuals each, and the population trend is stable there.[29]

Today, the northern subspecies occurs in West and Central Africa.[2] It is regionally extinct in Gambia, Mauritania, Sierra Leone, the Western Sahara, Morocco, Algeria, Tunisia, Libya and Egypt. Its range has declined to the:[39]

Former range

Dated to 1925, this is considered to be the last photograph of a wild North African lion

Historically, the northern subspecies' range encompassed North Africa.[2] In these regions, lion populations occurred in the Sahel, mountain ranges of the Sahara, Barbary Coast, Maghreb,[28][17][6] and the eastern Mediterranean Basin.[96][15] North African populations declined since the mid-19th century and were eradicated by the early 1960s.[101][6]

Characteristics

Adult male lions with long brown manes in Serengeti National Park, Tanzania
Male with a partially black mane in Narok County, southern Kenya
A mature male lion with an intermediate mane development in Amboseli National Park, Kenya
Young Serengeti male

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. Average head-to-body length of male lions is 2.47–2.84 m (8.1–9.3 ft) with a weight of 148.2–190.9 kg (327–421 lb). Females are smaller and less heavy.[96]

A few lion specimens from West Africa obtained by museums have been described as having shorter manes than lions from other African regions.[17] In general, the West African lion is similar in general appearance and size as lions in other parts of Africa and Asia.[18]

Zoological specimens range in colour from light to dark tawny. Male skins have short manes, light manes, dark manes or long manes.[102] Taxonomists recognised that neither skin nor mane colour and length of lions can be adduced as distinct subspecific characteristics. Then they turned to measuring and comparing lion skulls and found that skull length of Barbary and Indian lion samples does not differ significantly, ranging from 280–311.7 mm (11.02–12.27 in) in females and 338–362 mm (13.3–14.3 in) in males.[102][17]

A revision of lion skins in the

British Museum Natural History revealed that Central African lions are about the same size as Asiatic lions.[91]

The Cape lion had a black mane extending beyond the shoulders and under the belly.

Kalahari and eastern Okavango Delta alongside those with a normal tawny colour.[103] Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.[17][54]

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.[104]

Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa.[105] The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.[21]

Timbavati Private Game Reserve in South Africa. Their whitish fur is a rare morph caused by a double recessive allele.[106]

Manes

In the 19th and 20th centuries, lion type specimen were described on the basis of mane size and colour.

mane types. 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 800 m (2,600 ft) elevation 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.[108] Hence, lion manes reflect ambient temperature. The mane colour is also influenced by nutrition and testosterone. Its length is an indicator for age and fighting ability of the lion.[104]

A male lion specimen from Somalia had a short mane.[17] 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.[58] A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well.[24] 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.[57] Mane colour of males in Kenya vary between tawny, isabelline and light reddish yellow.[107] Tsavo male lions generally do not have a mane, though colouration and thickness vary. There are several hypotheses as to the reasons. One is that mane development is closely tied to climate because its presence significantly reduces heat loss.[109] An alternative explanation is that manelessness is an adaptation to the thorny vegetation of the Tsavo area in which a mane might hinder hunting. Tsavo males may have heightened levels of testosterone, which could also explain their reputation for aggression.[110]

The weak or absent mane of Tsavo lions is a feature which was characteristic also for the extinct lions of ancient Egypt and Nubia. Adult lion males in Egyptian art are usually depicted without a mane, but with a ruff around the neck.[111]

White lion

The white lion is a rare

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.[106] White lions are selected for breeding in captivity.[112] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[113]

Records

An exceptionally heavy male near

East African lion reportedly measured 3.33 m (10.9 ft).[96]

In 1936, a

Mucusso in southern Angola in 1973.[114][115]

Behaviour and ecology

The lion is a social cat, living in groups of related individuals with their offspring. Such a family group is called a 'pride'. The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have also been observed. Male lion groups are called a coalition. Membership only changes with the births and deaths of female lions. Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age.[116] The sole known exception of this pattern is the Tsavo lion pride, which always has just one adult male.[117]

Male lions spend years in a nomadic phase before gaining residence in a

radio-collared lions in the Kavango–Zambezi Transfrontier Conservation Area show that adult 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.[121]

Lions usually

cheetah, hyena and African wild dog.[116][118][123] The lion and spotted hyena occupy a similar ecological niche, and where they coexist, they compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%. Lions typically ignore spotted hyenas unless the lions are on a kill or are being harassed by the hyenas, while the latter tend to visibly react to the presence of lions, with or without the presence of food.[124]

Lions predominantly hunt large

prey is usually in the range of 40.0 to 270.0 kg (88.2 to 595.2 pounds).[122] Predation on adult African bush elephants has been observed in Chobe National Park, Botswana.[125][126]

Lions in Botswana's Okavango Delta have learned to swim in the delta's swamps. They hunt large prey like buffalo,[127] and occasionally also African elephants when smaller prey is scarce.[128]

In Benin's Pendjari National Park, groups of lions range from 1–8 individuals. Outside the national park, groups are smaller and more single male lions occur.[129]

Even within what was Sudan, the diet of lions varied, depending on availability of prey or geography. In the area of

man-eating were reported in the areas of the Bahr el-Ghazal and Atabarah.[81]

In what was Somaliland, lions were reported to have jumped over zaribas that were at least 10–12 ft (3.0–3.7 m) high to get even large livestock like camels out of them, or to consume them.[81]

Lions in Northeast Africa exhibit a variety of characteristics. Sudan lions, with the exception of one black-maned male near the Rahad River, were described as having scanty yellow manes. A desert-dwelling cub from Wadi Milh in West Kordofan was described as being pale, with the tail tuft and ear-backs being grey rather than black, and without clear spots.[81] A short-maned lion was also described from Somalia.[17]

Lions from the Ethiopian highlands were described as having manes that were dark and heavy.

captive lions at Addis Ababa Zoo.[21] Skulls of Ethiopian lions were shorter than those of Cape lions, which is a reason why authors like Heller considered the latter to have been bigger in body size.[81][130]

Attacks on humans

Threats

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.[39] 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.[75]

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.[142] Private game ranches in South Africa also breed lions for the canned hunting industry.[143]

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle.[144] 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.[145][146]

In 2015 and 2017, two male lions,

Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park.[147][148]

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.[149]

The West African lion is threatened by

illegal trade of body parts. Lion body parts from Benin are smuggled to Niger, Nigeria, Gabon, Ivory Coast, Senegal and Guinea, and from Burkina Faso to Benin, Ivory Coast, Senegal and Guinea.[150]

In Somalia's

trade them in markets, where they are priced between US$ 500 and 600 per cub. In the southern part of the country, cubs are also kept for export.[152]

Conservation

captive cubs at the Olusegun Obasanjo Library, Nigeria

African lions are included in

CITES Appendix II. Today, lion populations are stable only in large protected area complexes.[29]

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.[51][50] A significant incentive for local communities in a number of Southern African countries to support measures for conservation is that they generate significant revenue through wildlife tourism.[39]

In 2010, the small and isolated Kalahari population was estimated at 683 to 1,397 individuals in three protected areas, the

Gemsbok National Parks.[153] More than 2000 lions exist in the well-protected Kruger National Park.[154] In June 2015, seven lions were relocated from KwaZulu-Natal in South Africa to Akagera National Park in Rwanda.[155]

In 2006, a Lion Conservation Strategy for West and Central Africa was developed in cooperation between IUCN regional offices and several wildlife conservation organisations. 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.[156]

In captivity

Captive Kruger lion in Philadelphia Zoo

At the beginning of the 21st century, the

Haile Selassie I.[21][157]

In 2006, the registry of the

International Species Information System (ISIS) showed 29 lions that were derived from animals captured in Angola and Zimbabwe. In addition, about 100 captive lions were registered as P. l. krugeri by ISIS, which derived from lions captured in South Africa.[3][57] Interest in the Cape lion had led to attempts to conserve possible descendants in places like Tygerberg Zoo.[158][159]

In 2006, 1258

captive lions were registered in the International Species Information System, including 13 individuals originating from Senegal to Cameroon, and 970 with uncertain origin.[52]

  • Male in Khartoum Zoo, Sudan, 1936
    Male in Khartoum Zoo, Sudan, 1936
  • Major Barker, Director of Khartoum Zoo, playing with a young lioness in 1936. Note the rosettes.
    Major Barker, Director of Khartoum Zoo, playing with a young lioness in 1936. Note the rosettes.
  • Cubs at Addis Abeba Zoo
    Cubs at Addis Abeba Zoo

Cultural significance

Between 1910 and 1932, the Coat of arms of South Africa featured a lion
Coat of arms of Sierra Leone

The title "Lion of Mali" was given to Marijata of the Mali Empire.[160][161]

The Cameroon national football team is nicknamed "The Indomitable Lions."[162]

The lion is featured as an animal symbol in East Africa.[163][160] The name 'Simba' is a Swahili word for the lion, which also means 'aggressive', 'king' and 'strong'.[115]

The lion was important to the

mummified remains of a male lion in the tomb of Maia, the wet nurse of King Tutankhamun, on the opposite side of the Nile from Memphis. Nevertheless, lions were also hunted in large numbers by the Egyptians.[165] The ruins of Kushite city of Naqa in present-day Sudan include a Lion Temple.[166]

One of the lions that the Metro-Goldwyn-Mayer used as its mascot was Jackie.[167] It was brought from the Nubian part of Sudan.[168]

In December 2017, Sudan's President Omar al-Bashir had gifted 4 lion cubs to visiting Turkish President Recep Tayyip Erdoğan. Later housed at the Gaziantep Zoo, the cubs were seen as symbolising strength.[169]

Regional names

Lion populations in different parts of Africa were referred to by several regional names, including "Katanga lion", "Transvaal lion", "Kalahari lion",[24][59][60] "Southeast African lion", and "Southwest African lion",[90] in Southern Africa, "Masai lion", "Serengeti lion,"[116] "Tsavo lion"[110] and "Uganda lion"[17] in East Africa, and "Cameroon lion"[15] and "North-east Congo lion"[3] in Central Africa.

Gallery

See also

References

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Further reading

External links