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''Panthera leo melanochaita'' is a
lion The lion (''Panthera leo'') is a large cat of the genus '' Panthera'' native to Africa and India. It has a muscular, broad-chested body; short, rounded head; round ears; and a hairy tuft at the end of its tail. It is sexually dimorphic; adu ...
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
in
Southern Southern may refer to: Businesses * China Southern Airlines, airline based in Guangzhou, China * Southern Airways, defunct US airline * Southern Air, air cargo transportation company based in Norwalk, Connecticut, US * Southern Airways Express, M ...
and
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...
. In this part of
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, lion populations are regionally
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
in Lesotho,
Djibouti Djibouti, ar, جيبوتي ', french: link=no, Djibouti, so, Jabuuti officially the Republic of Djibouti, is a country in the Horn of Africa, bordered by Somalia to the south, Ethiopia to the southwest, Eritrea in the north, and the Red ...
and
Eritrea Eritrea ( ; ti, ኤርትራ, Ertra, ; ar, إرتريا, ʾIritriyā), officially the State of Eritrea, is a country in the Horn of Africa region of Eastern Africa, with its capital and largest city at Asmara. It is bordered by Ethiopi ...
, and are threatened by loss of habitat and prey base, killing by local people in retaliation for loss of livestock, and in several countries also by
trophy hunting Trophy hunting is a form of hunting for sport in which parts of the hunted wild animals are kept and displayed as trophies. The animal being targeted, known as the " game", is typically a mature male specimen from a popular species of collectab ...
. 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. In 2005, a Lion Conservation Strategy was developed for East and Southern Africa. Results of a phylogeographic study indicate that lion populations in southern and eastern Africa are forming a major
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
distinct from lion populations in West Africa, Central Africa and Asia. In 2017, the Cat Classification Task Force of the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
Cat Specialist Group subsumed lion populations according to the major clades into two subspecies, namely '' P. l. leo'' and ''P. l. melanochaita''. Within ''P. l. melanochaita'' three subclades are clearly distinguishable. One from northeastern Africa, another one from southwestern Africa and a third one from southeastern Africa. The
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
for ''P. l. melanochaita'' was a black-maned lion from the Cape of Good Hope, known as the Cape lion. Phylogeographic analysis of lion samples from
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the nort ...
and the
Republic of the Congo The Republic of the Congo (french: République du Congo, ln, Republíki ya Kongó), also known as Congo-Brazzaville, the Congo Republic or simply either Congo or the Congo, is a country located in the western coast of Central Africa to the w ...
indicate their close genetic relation to ''P. l. melanochaita'' samples from
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
and
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
. It has been referred to as the Southern lion, Southern African lion and the 'southern subspecies'.


Taxonomy

''Felis (Leo) melanochaitus'' was the
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bo ...
proposed by Charles Hamilton Smith in 1842 who described a lion specimen from South Africa's Cape Province. In the 19th and 20th centuries, several naturalists described
zoological specimen A zoological specimen is an animal or part of an animal preserved for scientific use. Various uses are: to verify the identity of a (species), to allow study, increase public knowledge of zoology. Zoological specimens are extremely diverse. Exampl ...
s from Southern and East Africa and proposed the following subspecies: *''Felis leo somaliensis'' (Noack 1891), based on two lion specimens from
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constitut ...
*''Felis leo massaicus'' (
Neumann Neumann is German and Yiddish for "new man", and one of the 20 most common German surnames. People * Von Neumann family, a Jewish Hungarian noble family A–G *Adam Neumann (born 1979), Israeli-born entrepreneur and founder of WeWork * Alfre ...
1900)
, based on two lions killed near Kibaya and the Gurui River in
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
*''Felis leo sabakiensis'' ( Lönnberg 1910), based on two male lions from the environs of
Mount Kilimanjaro Mount Kilimanjaro () is a dormant volcano in Tanzania. It has three volcanic cones: Kibo, Mawenzi, and Shira. It is the highest mountain in Africa and the highest free-standing mountain above sea level in the world: above sea level and a ...
*''Felis leo roosevelti'' ( Heller 1913), a lion from the Ethiopian Highlands presented to
Theodore Roosevelt Theodore Roosevelt Jr. ( ; October 27, 1858 – January 6, 1919), often referred to as Teddy or by his initials, T. R., was an American politician, statesman, soldier, conservationist, naturalist, historian, and writer who served as the 26t ...
*''Felis leo nyanzae'' (Heller 1913), a lion skin from
Kampala Kampala (, ) is the capital and largest city of Uganda. The city proper has a population of 1,680,000 and is divided into the five political divisions of Kampala Central Division, Kawempe Division, Makindye Division, Nakawa Division, and R ...
,
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
*''Felis leo bleyenberghi'' (Lönnberg 1914), a male lion from the Katanga Province of
Belgian Congo The Belgian Congo (french: Congo belge, ; nl, Belgisch-Congo) was a Belgian colony in Central Africa from 1908 until independence in 1960. The former colony adopted its present name, the Democratic Republic of the Congo (DRC), in 1964. Colo ...
*''Leo leo hollisteri'' ( Joel Asaph Allen 1924), a male lion from the area of
Lime Springs, Sotik Sotik Constituency is an Constituencies of Kenya, electoral constituency in Kenya established for the 1997 elections. It is one of five constituencies in Bomet County. Sotik has one major river, River Kipsonoi. Sotik is also a hilly place with the ...
on the eastern shore of
Lake Victoria Lake Victoria is one of the African Great Lakes. With a surface area of approximately , Lake Victoria is Africa's largest lake by area, the world's largest tropical lake, and the world's second-largest fresh water lake by surface area after ...
*''Leo leo krugeri'' ( Austin Roberts 1929), an adult male lion from the Sabi Sand Game Reserve named in honour of
Paul Kruger Stephanus Johannes Paulus Kruger (; 10 October 1825 – 14 July 1904) was a South African politician. He was one of the dominant political and military figures in 19th-century South Africa, and President of the South African Republic (or ...
*''Leo leo vernayi'' (Roberts 1948), a male lion from the Kalahari collected by the Vernay-Lang Kalahari Expedition *''Panthera leo webbensies'' Ludwig Zukowsky 1964, two lions from
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constitut ...
, one in the Natural History Museum, Vienna that originated in Webi Shabeelle, the other kept in a German zoo that had been imported from the hinterland of
Mogadishu Mogadishu (, also ; so, Muqdisho or ; ar, مقديشو ; it, Mogadiscio ), locally known as Xamar or Hamar, is the capital and List of cities in Somalia by population, most populous city of Somalia. The city has served as an important port ...
. Dispute over the validity of these purported subspecies continued among naturalists and curators of natural history museums until the early 21st century. In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies. In 1939, the American zoologist
Allen Allen, Allen's or Allens may refer to: Buildings * Allen Arena, an indoor arena at Lipscomb University in Nashville, Tennessee * Allen Center, a skyscraper complex in downtown Houston, Texas * Allen Fieldhouse, an indoor sports arena on the Univer ...
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. Pocock subordinated the lion to the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
'' Panthera'' in 1930, when he wrote about Asiatic lions. Ellerman and Morrison-Scott recognized only two lion subspecies in the
Palearctic realm The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Si ...
, namely the African ''P. l. leo'' and the Asiatic ''P. l. persica''. Various authors recognized between seven and 10 African lion subspecies. Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa. In the 1970s, the scientific name ''P. l. vernayi'' was considered synonymous with ''P. l. krugeri''. In 1975, Vratislav Mazák hypothesized that the Cape lion evolved geographically isolated from other populations by the
Great Escarpment The Great Escarpment is a major topographical feature in Africa that consists of steep slopes from the high central Southern African plateauAtlas of Southern Africa. (1984). p. 13. Reader's Digest Association, Cape Town downward in the directio ...
. 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 The Province of the Transvaal ( af, Provinsie van Transvaal), commonly referred to as the Transvaal (; ), was a province of South Africa from 1910 until 1994, when a new constitution subdivided it following the end of apartheid. The name "Trans ...
regions and farther east are considered having been possible through a corridor between the Great Escarpment and the
Indian ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by ...
. In 2005, the authors of '' Mammal Species of the World'' recognized ''P. l. bleyenberghi'', ''P. l. krugeri'', ''P. l. vernayi'', ''P. l. massaica'', ''P. l. hollisteri'' and ''P. l. nyanzae'' as valid taxa. In 2016,
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolo ...
assessors subsumed all African lion populations to ''P. l. leo''. Two lion subspecies are now recognised: *''P. l. melanochaita'' is understood as comprising lion populations in the contemporary Southern and East African range countries, * ''P. l. leo'' comprises lion populations in North, West and Central Africa and Asia.
Genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
-wide data of a wild-born historical lion sample from Sudan clustered with ''P. l. leo'' in mtDNA-based phylogenies, but with a high affinity to ''P. l. melanochaita''. This result indicates that the taxonomic position of lions in Central Africa may require revision.


Phylogeny

Since the beginning of the 21st century, several
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Scientists analysed between 32 and 480 lion samples from up to 22 countries. Results of
genetic analyses Genetic analysis is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts of ...
indicate that the
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
comprises two main
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary groups, one in Southern and East Africa, and the other in the northern and eastern parts of its historical range. These groups genetically diverged between 50,000 and 200,000 years ago. It was assumed that
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...
and the East African Rift constituted major barriers between the two groups. Lion samples from Gabon's
Batéké Plateau National Park Batéké Plateau National Park is a national park on the Bateke Plateau, southeastern Gabon covering . Due to its purported universal cultural and natural significance, it was added onto the UNESCO World Heritage Tentative List on October 20, ...
and
Odzala-Kokoua National Park Odzala-Kokoua National Park (or Odzala National Park) is a national park in the Republic of the Congo. The park was first protected in 1935, declared a biosphere reserve in 1977, and granted official designation by presidential decree in 2001. Od ...
in Republic of the Congo were found to be genetically closely related to lion samples from Namibia and Botswana. A phylogenetic analysis of lion samples from Africa and Asia showed that they shared a common ancestor probably between 98,000 and 52,000 years ago. Samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This indicates that Central Africa was a melting pot of lion populations after they had become isolated. They possibly migrated through corridors in the Nile Basin during the early
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
.


Overlap between subspecies

Among six samples of captive lions from Ethiopia, five clustered with samples from East Africa, but one with samples from the Sahel. For a subsequent phylogeographic study, eight wild lion samples from the Ethiopian Highlands were included in the
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...
analysis using 194 lion samples from 22 countries. Four of these samples
cluster may refer to: Science and technology Astronomy * Cluster (spacecraft), constellation of four European Space Agency spacecraft * Asteroid cluster, a small asteroid family * Cluster II (spacecraft), a European Space Agency mission to study th ...
ed with samples from Central Africa, and four with samples from East Africa, indicating that the
Great Rift Valley, Ethiopia The Great Rift Valley of Ethiopia, (or Main Ethiopian Rift or Ethiopian Rift Valley) is a branch of the East African Rift that runs through Ethiopia in a southwest direction from the Afar Triple Junction. In the past, it was seen as part of a " ...
was not a complete barrier to
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
. Southeastern Ethiopia is therefore considered a genetic admixture zone between Central and East African lions.


Distribution and habitat

In East and Southern Africa, lion populations declined in: *In
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
, where lion populations declined since at least the early 20th century due to trophy hunting by Europeans, killing of lions by local people out of fear, for illegal sale of skins and during civil wars. As of 2009, between seven and 23 lions were estimated to live in Nechisar National Park located in the Ethiopian Highlands. This small protected area is encroached by local people and their livestock. In 2012, lions were documented in cloud forest habitat of Kafa Biosphere Reserve. *
Somalia Somalia, , Osmanya script: 𐒈𐒝𐒑𐒛𐒐𐒘𐒕𐒖; ar, الصومال, aṣ-Ṣūmāl officially the Federal Republic of SomaliaThe ''Federal Republic of Somalia'' is the country's name per Article 1 of thProvisional Constitut ...
since the early 20th century. Intensive poaching since the 1980s and civil unrest posed a threat to lion persistence. *
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The ...
to near extinction in the 20th century. 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. Lions in Queen Elizabeth National Park form a contiguous population with lions in Virunga National Park. *
Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...
in the 1990s due to poisoning of lions and poaching of lion prey species. At least 108 lions were killed between 2001 and 2006 in the AmboseliTsavo WestTsavo East National Park network. As of 2006, there were an estimated 675 lions in the Tsavo national parks, out of the 2,000 total in Kenya. Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of . Lion populations in Kenya and Tanzania are fragmented over 17 patches ranging in size from . *
Rwanda Rwanda (; rw, u Rwanda ), officially the Republic of Rwanda, is a landlocked country in the Great Rift Valley of Central Africa, where the African Great Lakes region and Southeast Africa converge. Located a few degrees south of the Equator ...
and
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
due to killing of lions during the
Rwandan Civil War The Rwandan Civil War was a large-scale civil war in Rwanda which was fought between the Rwandan Armed Forces, representing the country's government, and the rebel Rwandan Patriotic Front (RPF) from 1October 1990 to 18 July 1994. The war aro ...
and ensuing
refugee crisis A refugee crisis can refer to difficulties and dangerous situations in the reception of large groups of forcibly displaced persons. These could be either internally displaced, refugees, asylum seekers or any other huge groups of migrants. A ...
in the 1990s. A small population was present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004. Lions were reintroduced to this national park in 2015. *
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...
and
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most central point. Its neighbours are t ...
due to illegal hunting of prey species in protected areas. *
Botswana Botswana (, ), officially the Republic of Botswana ( tn, Lefatshe la Botswana, label= Setswana, ), is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory being the Kal ...
due to intensive hunting and conversion of natural habitats for settlements since the early 19th century. In Northern Tuli Game Reserve, 19 lions died between 2005 and 2011 due to poaching, trophy hunting and
snaring Animal trapping, or simply trapping or gin, is the use of a device to remotely catch an animal. Animals may be trapped for a variety of purposes, including food, the fur trade, hunting, pest control, and wildlife management. History Neolithic ...
. *
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
due to massive killing of lions by farmers since at least the 1970s. In 2010, the small and isolated lion population in the Kalahari was estimated at 683 to 1,397 individuals in three protected areas, the Kgalagadi Transfrontier Park, the Kalahari Gemsbok and
Gemsbok National Park Kgalagadi Transfrontier Park is a large wildlife preserve and conservation area in southern Africa. The park straddles the border between South Africa and Botswana and comprises two adjoining national parks: * Kalahari Gemsbok National Park in S ...
s. *
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
since the early 19th century in the Natal and Cape Provinces south of the
Orange River The Orange River (from Afrikaans/Dutch: ''Oranjerivier'') is a river in Southern Africa. It is the longest river in South Africa. With a total length of , the Orange River Basin extends from Lesotho into South Africa and Namibia to the north ...
, where the Cape lion population was eradicated by 1860. A few decades later, lions in the Highveld north of the Orange River were also eradicated. In Transvaal, lions occurred historically in the Highveld as well, but were restricted to eastern Transvaal's
Bushveld The Bushveld (from af, bosveld, af, bos 'bush' and af, veld) is a sub-tropical woodland ecoregion of Southern Africa. It encompasses most of Limpopo Province and a small part of the North West Province of South Africa, the Central and Nort ...
by the 1970s. Between 2000 and 2004, 34 lions were reintroduced to eight protected areas in the Eastern Cape Province, including
Addo Elephant National Park Addo Elephant National Park is a diverse wildlife conservation park situated close to Gqeberha in South Africa and is one of the country's 20 national parks. It currently ranks third in size after Kruger National Park and the Kgalagadi Transfron ...
. In
Venetia Limpopo Nature Reserve {{Location map, South Africa Limpopo, float = right , width=250 , caption = Position of Venetia Limpopo Nature Reserve, label = Venetia Limpopo NR, position=background=white, lat=-22.30251, long=29.20412 The Venetia Limpopo Nature Reserve is ...
, 18 lions were trophy hunted and 11
euthanized Animal euthanasia (euthanasia from el, εὐθανασία; "good death") is the act of killing an animal or allowing it to die by withholding extreme medical measures. Reasons for euthanasia include incurable (and especially painful) conditio ...
between 2005 and 2011. Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped. Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals. The LCUs Ruaha−Rungwa, Serengeti−Mara, Tsavo−Mkomazi and Selous in East Africa, as well as Luangwa, Kgalagadi, Okavango−Hwange, Mid−Zambezi, Niassa and Greater Limpopo in Southern Africa are currently considered lion strongholds. These LCUs host more than 500 individuals each, and the population trend is stable there as of 2012.


North East African clade

Lions, which can be grouped into the northeastern clade are found in Somalia, Northern Kenya and Ethiopia, with a larger hybridization zone to the Northern lion subspecies (''P. l. leo'') in Ethiopia.


South East African clade

Lions, which can be grouped into the southeastern clade are found in Southern Kenya, Western
DRC The Democratic Republic of the Congo (french: République démocratique du Congo (RDC), colloquially "La RDC" ), informally Congo-Kinshasa, DR Congo, the DRC, the DROC, or the Congo, and formerly and also colloquially Zaire, is a country in ...
, Tanzania, Mozambique, Malawi, Zambia, southern Namibia and South Africa, with a larger hybridization zone to the southwestern lion clade in the Kruger National Park area (Great Limpopo Transfrontier Park). The classification of lions in Uganda is not clarified, since there were no samples included in the most comprehensive recent study.


South West African clade

Lions, which can be grouped into the southwestern clade are found in Angola, Northern Namibia, Northern Botswana and western Zimbabwe, extending southeast into the Tuli block. a There is a larger hybridization zone to the southeastern clade around the Kruger National Park.


Characteristics

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft.


Size and weight

Average head-to-body length of male lions is with a weight ranging from averaging in Southern Africa and averaging in East Africa. Females average in Southern Africa and in East Africa. Males in northern Kruger National Park weighed on average, whereas females weighed on average, and males in southern Kruger National Park weighed on average and females weighed , though there was an outbreak of
tuberculosis Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, ...
in southern park at the time. Skeletal muscles make up 58.8% of the lion's body weight. The largest known lion measured in length and weighed 375 kg (825 lb). An exceptionally heavy male lion near Mount Kenya weighed . The longest wild lion reportedly was a male shot near
Mucusso National Park Luengue-Luiana National Park ( pt, Parque Nacional do Luengue-Luiana) is a national park in Angola. Geography The park covers an area of 22,610 km². It located in Cuando Cubango Province in the southeastern corner of Angola. The park is bounde ...
in southern Angola in 1973. In 1936, a man-eating lion shot in 1936 in eastern Transvaal weighed about , and was considered to have been one of the heaviest wild lions. In 1963, two lions in Tanzania weighed after killing several
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to ani ...
.


Manes

In the 19th and 20th centuries, lion
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
s were described on the basis of their mane size and colour. Mane colour varies from sandy, tawny, isabelline, light reddish yellow to dark brown and black. Mane length varies from short to extending to knee joints and under the belly. Lions without a mane were observed in the Tsavo area. Mane development is related to age: older males have more extensive manes than younger ones; manes continue to grow up to the age of four to five years, long after lions become sexually mature. Males living in the Kenyan highlands above elevations of develop heavier manes than lions in the more humid and warmer lowlands of eastern and northern Kenya. Average
ambient temperature Colloquially, "room temperature" is a range of air temperatures that most people prefer for indoor settings. It feels comfortable to a person when they are wearing typical indoor clothing. Human comfort can extend beyond this range depending on ...
, nutrition and testosterone influence the colour and size of the mane. Its length is an indicator for age and fighting ability of the lion. In Serengeti National Park, female lions favour males with dense and dark manes as mates.


White lion

White lions have occasionally been encountered in and around South Africa's Kruger National Park and the adjacent Timbavati Private Game Reserve. Their whitish fur is a rare morph caused by a double
recessive In genetics, dominance is the phenomenon of one variant ( allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant an ...
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chrom ...
. It has normal pigmentation in eyes and skin. They were removed from the wild in the 1970s, thus decreasing the white lion
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
. Nevertheless, 17 births have been recorded in five different prides between 2007 and 2015. White lions were selected for breeding in
captivity Captivity, or being held captive, is a state wherein humans or other animals are confined to a particular space and prevented from leaving or moving freely. An example in humans is imprisonment. Prisoners of war are usually held in captivity by a ...
. They have been bred in South African camps for use as trophies to be killed during canned hunts.


Behaviour and ecology

In
Serengeti National Park The Serengeti National Park is a large national park in northern Tanzania that stretches over . It is located entirely in eastern Mara Region and north east portion of Simiyu Region and contains over of virgin savanna. The park was established ...
, monitoring of lion prides started in 1966. Between 1966 and 1972, two observed lion prides comprised between seven and 10 females each. Females had litters once in 23 months on average. Litters contained two to three cubs. Of 87 cubs born until 1970, only 12 reached the age of two years. Cubs died due to starvation in months when large prey was not available, or following take-over of the prides by new males. Male lions in coalitions are closely related. Between 1974 and 2012, 471 coalitions comprising 796 male lions entered a study area of . Of these, 35 coalitions included male lions that were born in the area but had left and returned after about two years of absence. Nomadic coalitions became resident at between 3.5 and 7.3 years of age. The lion population of Selous Game Reserve has been surveyed since 1996. Lion prides avoided
acacia ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus nam ...
woodlands and preferred habitats near water courses with short grasses, where also prey species gathered. Two or more prides shared home ranges. In Kavango–Zambezi Transfrontier Conservation Area, lions have been monitored since 1999. In 2003, 50 lions were radio-collared in Hwange National Park and tracked until 2012. Results show that adult male and female lions preferred grassland and
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It ...
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. In the semi-arid savanna of Zimbabwe's Hwange National Park, 19 lions were radio-collared and tracked between 2002 and 2007. Both female and male lions moved foremost within of waterholes in all seasons. Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002.


Hunting and diet

Lions usually hunt in groups and
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
foremost on ungulates such as
gemsbok The gemsbok or South African oryx (''Oryx gazella'') is a large antelope in the genus ''Oryx''. It is native to the extremely dry, arid regions of Southern Africa; notably, the Kalahari Desert. Some authorities formerly classified the East Afr ...
(''Oryx gazella''), Cape buffalo (''Syncerus caffer''),
blue wildebeest The blue wildebeest (''Connochaetes taurinus''), also called the common wildebeest, white-bearded gnu or brindled gnu, is a large antelope and one of the two species of wildebeest. It is placed in the genus '' Connochaetes'' and family Bovidae, a ...
(''Connochaetes taurinus''), giraffe (''Giraffa camelopardalis''), common eland (''Tragelaphus oryx''), greater kudu (''T. strepsiceros''),
nyala The lowland nyala or simply nyala (''Tragelaphus angasii'') is a spiral-horned antelope native to southern Africa. It is a species of the family Bovidae and genus ''Tragelaphus'', previously placed in genus ''Nyala''. It was first described in ...
(''T. angasii''), roan antelope (''Hippotragus equinus''), sable antelope (''H. niger''), plains zebra (''Equus quagga''), bushpig (''Potamochoerus larvatus''),
common warthog The common warthog (''Phacochoerus africanus'') is a wild member of the pig family (Suidae) found in grassland, savanna, and woodland in sub-Saharan Africa. In the past, it was commonly treated as a subspecies of ''P. aethiopicus'', but today th ...
(''Phacochoerus africanus''), hartebeest (''Alcephalus buselaphus''), common tsessebe (''Damaliscus lunatus''),
waterbuck The waterbuck (''Kobus ellipsiprymnus'') is a large antelope found widely in sub-Saharan Africa. It is placed in the genus '' Kobus'' of the family Bovidae. It was first described by Irish naturalist William Ogilby in 1833. Its 13 subspecies a ...
(''Kobus ellipsiprymnus''), kob (''K. kob'') and Thomson's gazelle (''Eudorcas thomsonii''). Their prey is usually in the range of . In the Serengeti National Park, lions were observed to also scavenge on carrion of animals that were killed by other predators, or died from natural causes. They kept a constant lookout for circling vultures, apparently being aware that vultures indicate a dead animal. Faeces of lions collected near waterholes in Hwange National Park also contained remains of climbing mice (''Dendromus'') and common mice (''Mus (genus), Mus''). In Botswana's Chobe National Park, lions also prey on young and subadult African bush elephants (''Loxodonta africana''). They successfully attacked 74 elephants between 1993 and 1996, of which 26 were older than nine years, and one bull over 15 years old. In October 2005, a pride of up to 30 lions killed eight African bush elephants that were between four and eleven years old.


Attacks on humans

Several cases of lion attacking people have been documented: *In the 1890s, two Tsavo Man-Eaters attacked workers during the building of the Uganda Railway. Their skulls and skins are part of the zoological collection of the Field Museum of Natural History. The total number of people killed is unclear, but allegedly 135 people fell victim to these lions in less than a year before John Henry Patterson (author), Colonel Patterson killed them. * The Njombe lions were a lion pride in Njombe in former Tanganyika (1961–1964), Tanganyika, which are thought to have preyed on 1,500 to 2,000 people. They were killed by George Gilman Rushby. *Between 1990 and 2004, lions killed more than 560 people in Tanzania, mostly during harvest season in crop fields and in areas where natural prey is scarce. * In February 2018, lions killed a suspected poacher near Kruger National Park. * In February 2018, Kevin Richardson (zookeeper), Kevin Richardson took three lions for a walk at Dinokeng Game Reserve in South Africa. A lioness pursued an impala for at least , and killed a young woman near her car. * In July 2018, human remains were found in the lion enclosure of a privately owned reserve in South Africa. They were suspected to have been rhino poachers, as they had a high-powered rifle with a silencer, an axe and wire cutters.


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. It is the primary cause for a decline of lion populations in Tanzania's Selous Game Reserve and Katavi National Park. 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. 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. Private game ranches in South Africa also breed lions for the canned hunting industry. In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle. 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. In 2015 and 2017, two male lions, Cecil (lion), Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park. Uncontrolled bushfires and hunting of lions and prey species in Zambia's Kafue National Park make it difficult for the lion population to recover. Cub mortality in particular is high.


Conservation

African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes.
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
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. Local communities in several Southern African lion range countries generate significant income through wildlife tourism, which is a strong incentive for their support of conservation measures. Establishing corridors between protected areas is important for facilitating dispersal of lions. Makgadikgadi Pans National Park and Central Kalahari Game Reserve are key dispersal areas in Southern Africa.


In captivity

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 of Ethiopia, Haile Selassie I. In 2006, eight captive lions were registered under the name ''P. l. massaicus'', and 23 as ''P. l. nubicus'' from Tanzania by the International Species Information System; about 100 captive lions were registered as ''P. l. krugeri'', which derived from lions captured in South Africa. In 2012, samples of lions kept at Sana'a Zoo in Yemen were found to cluster with those of lions from East and Southern Africa.


Cultural significance

The lion is an animal symbol in shamanistic rituals of the Nuer people. In other East African cultures, it symbolizes laziness. Scars inflicted by lions are regarded as a sign of courage among the Masai people. The name 'Simba' is a Swahili language, Swahili word for the lion, which also means 'aggressive', 'king' and 'strong'.


Regional names

Lion populations in Southern and East Africa were referred to by several regional names, including Katanga lion, Transvaal lion, Kalahari lion, Southeast African lion, and Southwest African lion, Masai lion, Serengeti lion, Tsavo lion and Uganda lion. It has also been referred to as 'Eastern-Southern African lion', 'Southern lion', and as 'southern subspecies'.


See also

*Wild cats in Africa: African leopard African golden cat Caracal Serval African wildcat Sand cat Black-footed cat Cheetah *''African Cats'' *''Born Free'' *Elsa the lioness *''The Lion King'' *Mapogo lion coalition *''Pride (2004 film), Pride''


References


External links

* *
$0.5m funding to stop the decline in the population of African lionsWhat Will It Take to Save the East African Lion from Extinction? Hunting or Herding?Lions in East Africa
*[https://petapixel.com/2018/09/04/lioness-steals-photographers-canon-dslr-and-gives-it-to-her-cubs/ PetaPixel, September 2018: Lioness Steals Photographer’s Canon DSLR and Gives It to Her Cubs]
National Geographic Wildlife Watch, June 2018: As Tigers Become Rarer, Poachers Are Targeting LionsLobengula the South African guardian lionWhat Happened to Angola’s 1,000 Lions?Holding the line for lions in MozambiqueGiant Lions Once Prowled East Africa, 200,000-Year-Old Skull RevealsBBC Earth: Lions take down an adult elephantThe Savuti Lions of the Chobe National ParkA Zambian lion stirsShamba the South African lionKnown for escapes, South African lion becomes a father
{{Taxonbar, from1=Q20907143, from2=Q2118614 Lions Subspecies Mammals described in 1842 Mammals of Angola Mammals of Botswana Mammals of the Democratic Republic of the Congo Mammals of Mozambique Mammals of Namibia Mammals of South Africa Mammals of Eswatini Mammals of Zimbabwe Mammals of Kenya Mammals of Tanzania Mammals of Ethiopia Mammals of Somalia Mammals of Uganda Fauna of East Africa Fauna of Southern Africa Mammals described in 1843 Mammals described in 1891 Mammals described in 1900