African lion

Male East African lion at Masai Mara, Kenya
Southern African lioness and cub near Otjiwarongo, Namibia
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 Horn of Africa or northern parts of East Africa is not fully resolved, due to the presence of both subspecies there, and lions in Central Africa have been grouped between the different subspecies by different genetic studies.[2][3] In addition, even the subspecies are not homogeneous,[1] being divided into different clades.[4]

Formerly described subspecies

Northwest Africa
SubspeciesDescription
Barbary lion (P. l. leo), also called the "Atlas lion", "Berber lion" or "North African lion"Formerly found in the Maghreb, this is 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
SubspeciesDescriptionImage
Senegal lion (P. l. senegalensis), also known as "West African lion"Found in West Africa.[10][11] It is listed as critically endangered as of 2015.

West Africa: Benin, Burkina Faso, Ghana,[12] Mali, Nigeria, Niger and Senegal

Gambian lion (P. l. gambianus)Formerly found in the Gambia.[13]
Central Africa
SubspeciesDescription
Cameroon lion (P. l. kamptzi)Found in Cameroon and the region south of Lake Chad, in Central or Western Africa.[14]
Northeast Congo lion (P. l. azandica), or simply the "Congo lion"Found in northeastern parts of the Congo, adjacent to Uganda.[10]

Central Africa:[15] (Democratic Republic of the Congo)

Northeast or East Africa
SubspeciesDescription
Nubian lion (P. l. nubica)From Nubia in Northeast Africa.[16]
Somali lion (P. l. somaliensis syn. P. l. webbiensisFrom Somaliland or Somalia, East Africa.[17][18]
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 Addis Ababa Zoo.[19] Researchers compared the microsatellite variations over ten loci of fifteen lions in captivity with those of six different wild lion populations. They determined that these lions are genetically unique and presumably that "their wild source population is similarly unique." These lions were part of a collection of the late Haile Selassie I of Ethiopia.[20]

Northeast Africa: Ethiopia

Kilimanjaro lion (P. l. sabakiensis)From the northern vicinity of Mount Kilimanjaro in East Africa.[21]
Ugandan lion (P. l. nyanzae)Found in Uganda, East Africa.[14]
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.[14]
Southern Africa
SubspeciesDescription
Cape lion (P. l. melanochaita)Formerly found from the Cape Province to Natal, South Africa.[22]
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.[17][23]

Central Africa: Formerly Katanga (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: (Botswana, Mozambique, South Africa, Swaziland, Zimbabwe)

Kalahari lion (P. l. vernayi)Found in the Kalahari Region of Southern Africa.[14]

Northern subspecies

Sultan the Barbary lion, New York Zoological Gardens, 1897

The Northern lion[24][25] (Panthera leo leo) is the nominate subspecies of the lion, which is present in the northern portion of Africa (particularly in West and Central Africa).[2][26] It is regionally extinct in North Africa. In West and Central Africa, lions are restricted to fragmented and isolated populations, most of them declining.[27][28] 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.[29] 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 phylogeographic study indicate that lion populations in West and Central African range countries are genetically close to populations in India, forming a clade distinct from lion populations in Southern and East Africa.[4] In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group subsumed lion populations to two subspecies, namely P. l. leo and P. l. melanochaita.[2]

Taxonomic history

A lion from Constantine, Algeria was the type specimen for the specific name Felis leo used by Linnaeus in 1758.[30] 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.[24][25] 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 and F. l. azandicus as valid taxa among ten lion subspecies.[22][16]
  • 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).[35]
  • Some authors considered P. l. nubicus a valid subspecies and synonymous with P. l. massaica.[16][17][36]
  • Some authors considered P. l. azandicus synonymous with P. l. massaicus and P. l. somaliensis, and P. l. kamptzi synonymous with P. l. senegalensis.[16][17]
  • In 2005, Wallace Christopher Wozencraft recognized P. l. kamptzi, P. l. bleyenberghi and P. l. azandica as valid taxa.[37]
  • In 2016, IUCN Red List assessors used P. l. leo for all African populations.[38]
  • 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 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 197 lion samples from up to 22 countries. They all agree that the lion 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.[39][40][41][42][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 hybrid lions. The hybrids had descended from lions captured in Angola and Zimbabwe, and apparently West or Central Africa. Results indicated that four lions from Morocco did not exhibit any unique genetic characteristics and shared mitochondrial haplotypes H5 and H6 with lions from West Africa, and together with them were part of a major 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[40]

Western and Central clades

Male West African lion at Pendjari National Park, Benin

Results of genetic analyses indicate that lions in West Africa and northern parts of Central Africa form distinct clades, which are more closely related to North African and Asiatic lions than to lions in Southern Africa and southern parts of East Africa. Lions from North Africa and India however, do form one single clade.[42] Analysis 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 Sahara from Senegal in the west to Nigeria in the east. This population has lost 99% of its former range.[43] It is regionally extinct in Mauritania, Gambia, Guinea-Bissau, Mali, Sierra Leone, Ivory Coast, and Togo, and possibly extinct in Guinea.[29] 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.[43][44]

The range of the Central African lion clade reaches from the lower Niger river in West Africa to Ethiopia, encompassing Cameroon, Central African Republic, northern parts of the Democratic Republic of the Congo, southern parts of Chad and Sudan, and South Sudan.[4] Some Central African populations permanently inhabit rainforests and clearings in rainforest mixed with savannah grassland.[27] In Cameroon, lions are present in Bénoué National Park, and smaller lion groups also in Waza National Park.[26] Habitat in Waza National Park comprises foremost dry woodland that is partly flooded during the rainy season from July to December. In this protected area, two radio-collared male lions used home ranges of between 428 and 1,054 km2 (165 and 407 sq mi), both inside and outside the park. Three radio-collared females had home ranges of between 352 and 724 km2 (136 and 280 sq mi) and stayed inside the park during most of the survey period.[45] In the Central African Republic, lions are present in Bamingui-Bangoran National Park and Biosphere Reserve, Manovo-Gounda St. Floris and Awakaba National Parks, Aouk Aoukale, Yata Ngaya, Nana Barya and Zemongo Faunal Reserves, and in several hunting reserves of the country.[46] Estimated lion numbers in the country are generally thought to be unreliable.[26][28] Lions in Virunga National Park form a contiguous population with lions in the East African country of Uganda.[26][28] In Chad, lions inhabit 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.[27] In 2004, the lion population in the country was estimated at maximum 225 individuals.[26] The following table shows estimates of lion population sizes between 2002 and 2012:

Range countriesArea used in km2Estimated no. of individuals
Bénoué National Park complex, Cameroon14,682200[47]
Waza National Park in Cameroon1,45217[48]
Central African Republic339,418about 1,297[46]
Garamba National Park and Domaine Chasse Bili Uere, Democratic Republic of the Congo115,671175−320[28][27]
Southeastern Chad133,408400[28]
Southwestern Sudan331,834375[28]
Boma and Gambela National Parks, South Sudan and Ethiopia106,941500[28]
Omo National Park, Sudan and Ethiopia22,483200[28]
other protected and non-protected areas, Ethiopia70,75996[28]
Total1,136,6483,260−3,305

Contemporary lion distribution and habitat quality in savannahs of West and Central Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[49] Educated guesses for size of populations in these LCUs ranged from 3,274 to 3,909 individuals between 2002 and 2012.[27][28]

Range countriesLion Conservation UnitsArea in km2
Senegal, Mali, Guinea-Bissau, GuineaNiokolo-Koba National Park90,384[28]
GuineaNational Park of Upper Niger613[28]
Benin, Burkina Faso, NigerW-Arly-Pendjari Complex29,403[28]
Beninthree unprotected areas6,833[28]
NigeriaYankari National Park and Kainji National Park6,551[28]
CameroonWaza and Bénoué National Parks16,134[48]
Central African Republiceastern part of the country; Bozoum and Nana Barya Faunal Reserves339,481[46]
Chadsoutheastern part133,408[28]
Democratic Republic of CongoGaramba-Bili Uere115,671[49]
South Sudan, Sudan331,834[49]
South Sudan, EthiopiaBoma-Gambella106,941
EthiopiaSouth Omo, Nechisar, Bale, Welmel-Genale, Awash National Parks, Ogaden93,274[28]

Southern subspecies

Lions shot in Kenya's Sotik Plains in 1909. Sotik lions had been recognized as a subspecies, with the scientific name Panthera leo hollisteri.[37][18]

The Southern lion (Panthera leo melanochaita),[24][25] also referred to as the East-Southern African lion[50] or Eastern-Southern African lion,[3] is a subspecies of the lion in Southern and East Africa.[2][51] In this part of Africa, lion populations are regionally extinct in Lesotho, Djibouti and Eritrea.[38] 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.[52] 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.[38]

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.[53]

Taxonomic history

Charles Hamilton Smith described the type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus.[54] It was referred to as the "Southern relative" of the North African lion.[24][25] 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.[53][37][22][61][16]

In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies.[58][22][61][62] 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.[22]

Pocock subordinated lions to the genus Panthera in 1930, when he wrote about Asiatic lions.[63] 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.[64] Various authors recognized between seven and 10 African lion subspecies.[16] Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.[65]

In the 1970s, the scientific name P. l. vernayi was considered synonymous with P. l. krugeri.[16] In 1975, Vratislav Mazák hypothesized that the Cape lion evolved geographically isolated from other populations by the Great Escarpment.[53] 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.[3][66]

In 2005, the authors of Mammal Species of the World recognized P. l. bleyenberghi and P. l. krugeri, P. l. vernayi P. l. massaica, P. l. hollisteri and P. l. nyanzae as valid taxa.[37] In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo.[38] 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

The Cape lion, the type specimen for the southern subspecies (P. l. melanochaita). Genetic research has shown it to be closely related to extant Southern African lions.[66]

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 197 lion samples from up to 22 countries. Based on the results of a genetic analyses, it appears that the species comprises two main evolutionary groups, one in Southern and East Africa, and the other in the northern and eastern parts of its historical range; these groups diverged about 50,000 years ago.[67] It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.[3][40][68][42][4] Based on this assessment, the species comprises two recognised subspecies:[2]

  • P. l. leo in the northern and eastern regions of the species' historical and contemporary distribution
  • P. l. melanochaita in Southern and East African range countries.

The two groups were in contact in Ethiopia or northern parts of East Africa.[4] A phylogeographic analysis of 194 lion sequences from 22 countries indicated that East African and Southern African lions form a clade that diverged about 186,000–128,000 years ago from the clade formed by North, 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 phylogeographic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Results of a DNA analysis using 26 lion samples from Southern and East Africa indicate that genetic variation between them is low and that two major clades exist: one in southwestern Africa and one in the region from Uganda and Kenya to KwaZulu-Natal. Five lion samples from Kenya's Tsavo East National Park showed identical haplotypes as three lion samples from the Transvaal region in South Africa.[69] 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.[40][42] 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.[40]

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.[20]

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.[70] In northern Kenya, lions had been observed near Kavirondo, near Lake Manyara and in the Tanga Region in the late 19th century.[56] By the 21st century, lion populations in northern Kenya have been fragmented.[71]

Range countriesArea used in km2Estimated no. of individuals
Laikipia-Samburu complex in Kenya35,511271[28]
Meru in Kenya7,36540[28]
Arawale complex in Kenya and Somalia22,540750[28]
Arboweerow-Alafuuto in Somalia24,527175[28]
TotalXXX XXX

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:[28]

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,[28][72][26] 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.[73] As of 2006, there were an estimated 675 lions in the Tsavo area, out of the 2,000 total in Kenya.[74] Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of 3,684 km2 (1,422 sq mi).[71]

A small population is present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004.[26]

The population in South Africa's former Natal and Cape Provinces is locally extinct since the mid 19th century.[75] The last lions south of the Orange River were sighted between 1850 and 1858.[53] Between 2000 and 2004, 34 lions were reintroduced to eight protected areas in the Eastern Cape Province, including Addo Elephant National Park.

Range countriesArea used in km2Estimated no. of individuals
Virunga and Queen Elizabeth National Park in CAR and Uganda5,583210[28]
Lake Mburo in Uganda3733[28]
Luama Hunting Reserve in DRC5,197<50[28]
Itombwe Massif in DRC3,244<50[28]
North West Tansania4,703105[28]
Ruaha-Rungwa in Tanzania195,9933,779[28]
Mpanga Kipengere in Tanzania95814[28]
Swaga Swaga in Tanzania7,242102[28]
Serengeti-Mara in Tanzania and Kenya35,8523,673[28]
Nairobi in Kenya830<30[28]
Tsavo-Mkomazi in Kenya and Tanzania39,216880[28]
Tarangire In Tanzania28,771731[28]
Wami Mbiki-Saadani in Tanzania8,787136[28]
Selous in Tanzania138,0357,644[28]
Niassa in Mozambique, Tanzania177,5591,573[28]
Liuwa Plains in Zambia3,8664[28]
Kafue in Zambia58,898386[28]
Nsumbu in Zambia5,650<50[28]
Luangwa National Park in Zambia72,992574[28]
Kasungu in Malawi2,3414[28]
Nkhotakota in Malawi1,84618[28]
Kgalagadi in South Africa and Botswana163,329800[28]
Mid-Zambezi in Mozambique, Zambia, Zimbabwe and Mozambique64,672755[28]
Tete South of Cahora Bassa, Gile and Gorongosa-Marromeu in Mozambique13,612, 22,322, 46,78159, 45, 229[28]

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.[28]

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[76][28] Another important reserve for this clade is the Etosha National Park.[28] Lions are considered regionally extinct in the southwestern part of the Democratic Republic of the Congo.[38][5]

In Gabon, the presence of lions in Batéké Plateau National Park was doubtful in 2010.[77] In 2015, a camera trap recorded a single male lion in this national park.[78] Continued camera trapping in the area for more than one year recorded the same lion repeatedly. Its hair samples were collected for 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.[79]

Range countriesArea used in km2Estimated no. of individuals
Southwestern clade
Kissama-Mumbondo in Angola4,593<10[28]
Bocoio-Camucuio in Angola22,00555[28]
SE Angola386,9621,905[28]
Sioma Ngwezi in Zambia4,155<50[28]
Etosha-Kunene in Namibia123,800455[28]
Khaudum-Caprivi in Namibia92,372150[28]
Xaixai in Botswana12,48475[28]
Okavango-Hwange99,5522,300[28]
Greater Mapungubwe in Botswana, South Africa and Zimbabwe5,15825[28]
Bubye in Zimbabwe4,875200[28]
TotalXXX XXX

Northeast African population

Male at Addis Ababa Zoo, Ethiopia

In Northeast Africa, lions occur in Ethiopia, Somalia, South Sudan and Sudan, but are regionally extinct in Djibouti and Eritrea.[38] The results of genetic tests published in 2016 indicated that lions in the region where North Africa overlaps with East Africa (including northern Kenya and possibly northern Uganda), are genetically mixed[4] between the Central African (P. l. leo) and East-Southern African lions (P. l. melanochaita). As a consequence, the taxonomic status of lions in the Horn of Africa was not resolved by the Cat Specialist Group.[2] Subpopulations were referred to by 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",[80] depending on the locality of occurrence.[17][14][81]

Genetic analyses and taxonomic history

Young male Nubian lion in New York Zoo, 1903[82]
Abyssinian lion in New York Zoo, 1914[83]

In the 19th century, a number of subspecies were described for lions in Northeast Africa. For example, zoological specimens from Nubia and Somalia were described or proposed by zoologists under the trinomina Felis leo nubicus[32] and Felis leo somaliensis.[55] In later centuries, these trinomina were alternatively considered to be synonymous with the scientific names of the North[22][37] and East African lions.[17][81]

A test done in 2012 on 15 lions at Addis Ababa Zoo and lions from 6 wild populations demonstrated that the captive lions were genetically different to wild lions in other parts of East Africa, but similar to wild lions from Cameroon and Chad.[20][19]

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.[42] 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 phylogeographic analysis using samples from African and Asiatic lions was published in 2006. One of the African samples was a vertebra from the 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 Pleistocene cave lions, the results of which were published in 2009, revealed that a lion sample from Sudan was distinct from lion samples that originated in the northeastern part of Congo-Kinshasa.[84]

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 Gambella. According to genetic research, this population, which is contigous with populations in Sudan, does not belong to the Southern subspecies but to the Northern lion. The same is probably true for the populations in northern Ethiopia,[4] where, a group of lions was recorded in 2016 in Alatash National Park close to the international border with Sudan.[85][86][87]

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.[26] 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.[88]

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.[26][28]

Central African population

A Northeast Northeast Congolese lioness (formerly P. l. azandica)[18][89] with a partial mane in Virunga National Park, which is adjacent to Queen Elizabeth National Park in Uganda. The Southeast African lion (P. l. melanochaita)[2] is believed to be in parts of D. R. Congo or Central Africa that are adjacent to East African countries like Uganda.[4]
Captive Cameroon lions (formerly P. l. kamptzi)[34][14] at Mefou National Park, Central Africa. Cameroon lions belong to the Central clade of the northern subspecies[4] (P. l. leo).[2]

The Central African lion[90] is a population of lions in Central Africa that has been grouped under the northern subspecies (Panthera leo leo), but was also found to be related to the southern subspecies[3][4] (Panthera leo melanochaita),[2][38] depending on the subpopulation, and is fragmented into small and isolated groups since the 1950s.[27][26]

Distribution and habitat

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

In Northeast Africa, lions may inhabit a variety of habitats, excepting dense forests. For example, in Sudan, they were recorded on the banks of the Blue and White Niles, and far away into desert areas.[80]

Currently, not much is known about the distribution and sizes of populations in Sudan]] and South Sudan. In the 1980s, lions were reported in Southern Kordofan and Southern Darfur provinces, located west of the Nile River.[27][28] In 2016, lions were discovered in Ethiopia's Alatash National Park, which is adjacent to Sudan's Dinder National Park.[85][86][87]

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.[91]
  • Uganda to near extinction in the 20th century.[92]
  • Kenya in the 1990s due to poisoning of lions and poaching of lion prey species.[27]
  • Rwanda and Tanzania due to killing of lions during the Rwandan Civil War and ensuing refugee crisis in the 1990s.[27]
  • Malawi and Zambia due to illegal hunting of prey species in protected areas.[27]
  • Botswana due to intensive hunting and conversion of natural habitats for settlements since the early 19th century.[93]
  • Namibia due to massive killing of lions by farmers since at least the 1970s.[94]
  • 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.[53] A few decades later, lions in the Highveld north of the Orange River were also eradicated.[95] In the Transvaal, lions occurred historically in the Highveld as well, but were restricted to eastern Transvaal's Bushveld by the 1970s.[96]

Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[50] Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals.[27][28]

Range countriesLion Conservation UnitsArea in km2
Democratic Republic of CongoMassif D'itombwe, Luama8,441[50]
Democratic Republic of Congo, UgandaQueen Elizabeth-Virunga5,583[97]
UgandaToro-Semulik, Lake Mburo, Murchison Falls4,800[98]
SomaliaArboweerow-Alafuuto24,527[50]
Somalia, KenyaBushbush-Arawale22,540[50]
KenyaLaikipia-Samburu, Meru and Nairobi National Parks43,706[28]
Kenya, TanzaniaSerengeti-Mara and Tsavo-Mkomazi75,068[46]
TanzaniaDar-Biharamulo, Ruaha-Rungwa, Mpanga-Kipengere, Tarangire, Wami Mbiki-Saadani, Selous384,489[46]
Tanzania, MozambiqueNiassa177,559[99]
MozambiqueCahora Bassa, Gilé, Gorongosa-Marromeu82,715[99]
Mozambique, ZambiaMiddle Zambezi64,672[99]
Mozambique, South AfricaGreat Limpopo Transfrontier Park150,347[99]
ZambiaLiuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex72,569[28]
Zambia, MalawiNorth-South Luangwa72,992[28]
MalawiKasungu, Nkhotakota4,187[28]
ZimbabweMapungubwe, Bubye10,033[28]
Botswana, ZimbabweOkavango-Hwange99,552[28]
BotswanaXaixai12,484[50]
Botswana, South AfricaKgalagadi163,329[28]
AngolaKissama-Mumbondo, Bocoio-Camacuio, Alto Zambeze393,760[50]
Angola, NamibiaEtosha-Kunene123,800[50]
NamibiaKhaudum-Caprivi92,372[50]

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.[28]

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:[38]

  • West African lion population surviving in a few protected areas of Senegal, Burkina Faso, Niger, Benin and Nigeria.[29] This population is fragmented and isolated, comprising fewer than 250 mature individuals.[29] The Central African population is threatened by loss of habitat and prey base and trophy hunting.[27][48][47]
  • Central African lion population surviving in protected areas of Cameroon, the Central African Republic, northern parts of the Democratic Republic of the Congo, southern parts of Chad and South Sudan and in Ethiopia.[28][47][48][46]

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,[27][16][6] and the eastern Mediterranean Basin.[95][14] North African populations declined since the mid-19th century and were eradicated by the early 1960s.[100][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

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.[95]

A few lion specimens from West Africa obtained by museums have been described as having shorter manes than lions from other African regions.[16] In general, the West African lion is similar in general appearance and size as lions in other parts of Africa and Asia.[17]

Zoological specimens range in colour from light to dark tawny. Male skins have short manes, light manes, dark manes or long manes.[101] 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.[101][16]

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 (97–112 in) with a weight of 148.2–190.9 kg (327–421 lb). Females are smaller and less heavy.[95]

The Cape lion had a black mane extending beyond the shoulders and under the belly.[54] Yet, black-maned lions also occur in the Kalahari and eastern Okavango Delta alongside those with a normal tawny colour.[102] Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.[16][53]

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.[103]

Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa.[104] The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.[20]

White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in South Africa. Their whitish fur is a rare morph caused by a double recessive allele.[105]

Manes

In the 19th and 20th centuries, lion type specimen were described on the basis of mane size and colour.[106] Male East African lions are known for a great range of 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.[107] 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.[103]

A male lion specimen from Somalia had a short mane.[16] 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.[57] A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well.[23] 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.[56] Mane colour of males in Kenya vary between tawny, isabelline and light reddish yellow.[106] 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.[108] 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.[109]

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.[110]

White lion

White lions owe their coloring to a recessive gene[105]

The white lion is a rare morph of the South African lion with a genetic condition called leucism, which is caused by a double recessive allele. It has normal pigmentation in eyes and skin. White individuals have been occasionally encountered only in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five different prides between 2007 and 2015.[105] White lions are selected for breeding in captivity.[111] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[112]

Records

An exceptionally heavy male near Mount Kenya weighed 272 kg (600 lb).[5] The largest East African lion reportedly measured 3.33 m (10.9 ft).[95]

In 1936, a man-eating South African lion shot by Lennox Anderson, outside Hectorspruit in Eastern Transvaal, weighed about 313 kg (690 lb), and was considered to be the heaviest wild lion. The longest wild lion reportedly was a male shot near Mucusso in southern Angola in 1973.[113][114]

Behaviour and ecology

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

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.[115] The sole known exception of this pattern is the Tsavo lion pride, which always has just one adult male.[116]

Male lions spend years in a nomadic phase before gaining residence in a pride.[117] In 1966, a program was started to monitor lions in Serengeti National Park.[115] Between 1966 and 1972, two observed prides had between seven and ten females each. On average, females had litters once in 23 months.[118] Two or three cubs comprised the litters, and only twelve managed to grow to the age of two, out of 87 cubs born until 1970. Cubs died due to starvation in months Factors that contributed to the deaths of cubs were starvation, when large prey was not available, or when new males took over the prides. Between 1974 and 2015, prides were monitored again, and until 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 this place but had left and returned after being absent for about two years. Nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[119] Results of a 10-year long survey on 50 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.[120]

Lions usually hunt in groups and prey foremost on ungulates such as wildebeest, zebra, African buffalo, gemsbok and giraffe.[121] In the Serengeti National Park, lions were observed to also scavenge on carrion when the opportunity arises. They scavenged 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. Sympatric predators include the leopard, cheetah, hyena and African wild dog.[115][117][122] 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.[123]

Lions predominantly hunt large ungulates like zebra, warthog, blue wildebeest, impala, gemsbok, Thomson's gazelle, kob, waterbuck, kudu, giraffe and Cape buffalo. Their prey is usually in the range of 40.0 to 270.0 kg (88.2 to 595.2 pounds).[121] Predation on adult African bush elephants has been observed in Chobe National Park, Botswana.[124][125]

Lions in Botswana's Okavango Delta have learned to swim in the delta's swamps. They hunt large prey like buffalo,[126] and occasionally also African elephants when smaller prey is scarce.[127]

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.[128]

Even within what was Sudan, the diet of lions varied, depending on availability of prey or geography. In the area of Lake No and the Rivers of Zeraf and Bahr el-Ghazal, they preyed on white-eared kobs. On the banks of the Atbara, Rahad, Blue Nile and lower White Nile, they tended to take livestock. In the eastern part of the country, camels were taken. In desert areas far away from sources of water, like in West Kordofan, lions were reported to consume watermelons for obtaining moisture, as was the case with jackals and people. A few cases on man-eating were reported in the areas of the Bahr el-Ghazal and Atabarah.[80]

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.[80]

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.[80] A short-maned lion was also described from Somalia.[16]

Lions from the Ethiopian highlands were described as having manes that were dark and heavy.[129] A similar observation was made for the manes of captive lions at Addis Ababa Zoo.[20] 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.[80][129]

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.[38] 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.[74]

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

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle.[143] 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.[144][145]

In 2015 and 2017, two male lions, Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park.[146][147]

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.[148]

The West African lion is threatened by poaching and 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.[149]

In Somalia's El Buur District,[70][150] intensive poaching since the 1980s and civil unrest posed a threat to lion persistence.[27] In the Lower Shabeelle, lionesses are killed by hunters who collect cubs to 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.[151]

Conservation

Two 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.[28]

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.[50][49] 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.[38]

In 2010, the small and isolated Kalahari population was estimated at 683 to 1,397 individuals in three protected areas, the Kgalagadi Transfrontier Park, the Kalahari Gemsbok and Gemsbok National Parks.[152] More than 2000 lions exist in the well-protected Kruger National Park.[153] In June 2015, seven lions were relocated from KwaZulu-Natal in South Africa to Akagera National Park in Rwanda.[154]

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.[155]

In captivity

Captive Kruger lion in Philadelphia Zoo

At the beginning of the 21st century, the Addis Ababa Zoo kept 16 adult lions. It is assumed that their ancestors, five males and two females, were caught in southwestern Ethiopia as part of a zoological collection for Emperor Haile Selassie I.[20][156]

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][56] Interest in the Cape lion had led to attempts to conserve possible descendants in places like Tygerberg Zoo.[157][158]

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.[51]

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.[159][160]

The Cameroon national football team is nicknamed "The Indomitable Lions."[161]

The lion is featured as an animal symbol in East Africa.[162][159] The name 'Simba' is a Swahili word for the lion, which also means 'aggressive', 'king' and 'strong'.[114]

The lion was important to the ancient Egyptians and Nubians. One of the defining pieces of Ancient Egyptian architecture is the Great Sphinx of Giza, the body of which resembles that of a lion, barring the human head.[163] In 2004, a team of archeologists under the leadership of Alain Zivie found 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.[164] The ruins of Kushite city of Naqa in present-day Sudan include a Lion Temple.[165]

One of the lions that the Metro-Goldwyn-Mayer used as its mascot was Jackie.[166] It was brought from the Nubian part of Sudan.[167]

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.[168]

Regional names

Lion populations in different parts of Africa were referred to by several regional names, including "Katanga lion", "Transvaal lion", "Kalahari lion",[23][58][59] "Southeast African lion", and "Southwest African lion",[89] in Southern Africa, "Masai lion", "Serengeti lion,"[115] "Tsavo lion"[109] and "Uganda lion"[16] in East Africa, and "Cameroon lion"[14] and "North-east Congo lion"[3] in Central Africa.

See also

References

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Further reading

  • The Penny Cyclopædia of the Society for the Diffusion of Useful Knowledge. 14. Charles Knight and Co. 1846-01-09. Retrieved 2014-08-28.
  • Charles Knight, ed. (1867). The English Cyclopaedia. Retrieved 2014-08-28.
  • Beolens, B.; Watkins, M.; Grayson, M. (2009-10-07). The Eponym Dictionary of Mammals. Baltimore: Johns Hopkins University Press. p. 110. ISBN 978-0-8018-9533-3.
  • Funaioli, U; Simonetta, A. M. (1966). "The Mammalian Fauna of the Somali Republic: Status and Conservation Problems". The Mammalian Fauna of the Somali Republic: Status and Conservation Problems. 1 (The Mammalian Fauna of the Somali Republic: Status and Conservation Problems). doi:10.1080/03749444.1966.10736746 (inactive 2018-09-27). Retrieved 19 September 2018.
  • Kays, Roland W.; Patterson, Bruce D. (2002). "Mane variation in African lions and its social correlates". Canadian Journal of Zoology. 80 (3): 471–478. doi:10.1139/z02-024. ISSN 0008-4301.
  • Schofield, A. (2013). White Lion: Back to the Wild. Pennsauken: BookBaby. ISBN 978-0620570053.
  • Selous, F. C. (2011). "XXV". Travel and Adventure in South-East Africa. New York: Cambridge University Press. p. 445. ISBN 978-1108031165.
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