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Zoological name: Panthera leo melanochaita
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The Southern lion (Panthera leo melanochaita) is a lion subspecies in Southern and East Africa. In this part of Africa, lion populations are regionally extinct in Lesotho, Djibouti and Eritrea. Since the turn of the century, lion populations in intensively managed protected areas in Botswana, Namibia, South Africa and Zimbabwe have increased, but declined in East African range countries.
The type specimen for P. l. melanochaita was a black-maned lion from the Cape of Good Hope, known as the Cape lion. The lion population in this part of South Africa is extinct. Living lion populations in other parts of Southern Africa were treated as distinct subspecies and referred to by several regional and scientific names, including "Katanga lion" (P. l. bleyenberghi), "Transvaal lion" (P. l. krugeri), "Kalahari lion" (P. l. vernayi), "Southeast African lion", and "Southwest African lion". Likewise, populations in East Africa were treated as distinct subspecies or referred to by different names, such as "Masai lion" (P. l. massaica), "Serengeti lion," "Tsavo lion", and "Uganda lion" (P. l. nyanzae), depending on the countries where individuals had been captured for zoological collections.
In the 19th and 20th centuries, several lion type specimens from Southern Africa were described and proposed as subspecies:
- In 1842, Charles Hamilton Smith described a black-maned lion from the Cape of Good Hope under the name Felis (Leo) melanochaitus. Naturalists and hunters of the 19th century recognized it as a distinct subspecies because of its dark mane colour.
- In 1891, two lions caught in Somalia were described under Felis leo somaliensis.
- In the 1890s, the German zoologist Neumann observed lions in East Africa. He proposed the trinomen Felis leo massaicus based on two type specimens, one male killed near Kibaya and one female killed at the Gurui River. They differ in mane size from lions caught in Somalia.
- In 1900, the Swedish zoologist Lönnberg described two lion specimens from the environs of Mount Kilimanjaro using the name Felis leo sabakiensis that were killed during a Swedish zoological expedition to East Africa.
- In 1914, Lönnberg described a male lion collected in the Katanga Province of Belgian Congo as type specimen of Felis leo bleyenberghi.
- In 1914, the American zoologist Heller described a lion from the Ethiopian Highlands under the name Felis leo roosevelti based on a male lion presented to Theodore Roosevelt. He also described a lion skin from Kampala under the name Felis leo nyanzae that differed slightly in skin and mane colour from F. l. massaica.
- In 1924, Joel Asaph Allen described a male lion as type specimen for Leo leo hollisteri that originated near Lime Springs, Sotik on the eastern shore of Lake Victoria and was obtained by the American Museum of Natural History.
- In 1929, Austin Roberts described an adult male lion with a dark brown mane from the Sabi Sand Game Reserve as type specimen for the Kruger lion Leo leo krugeri, named in honour of Paul Kruger.
- In 1948, Roberts also described a yellow-maned male lion specimen from the Kalahari as type specimen for the Kalahari lion Leo leo vernayi. It had been collected by the Vernay-Lang Kalahari Expedition in 1930.
- In 1964, the German zoologist Ludwig Zukowsky described two lions from Somalia as P. l. webbensies. One was a mounted specimen in the Natural History Museum, Vienna that originated in Webi Shabeelle, the other an individual kept in a German zoo that had been imported from the hinterland of Mogadishu.
In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies. 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.
Pocock subordinated lions to the genus Panthera in 1930, when he wrote about Asiatic lions. 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. 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. 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.
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. In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo. In 2017, the Cat Classification Task Force of the Cat Specialist Group reduced the number of valid lion subspecies in Southern and East Africa to one, namely P. l. melanochaita.
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. 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. 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.
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.
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 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 Africa lions.
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). The largest East African lion measured 3.33 m (10.9 ft). Females are smaller and less heavy.
Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa. The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.
In the 19th and 20th centuries, lion type specimen were described on the basis of mane size and colour. 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. 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.
A male lion specimen from Somalia had a short mane. 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. A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well. 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. Mane colour of males in Kenya vary between tawny, isabelline and light reddish yellow. 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. 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.
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.
The white lion is a rare morph 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. White lions are selected for breeding in captivity. Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.
In 1936, a man-eating lion shot by Lennox Anderson, outside Hectorspruit in Eastern Transvaal weighed about 313 kg (690 lb) and was the heaviest wild lion on record. The longest wild lion reportedly was a male shot near Mucusso in southern Angola in 1973.
Distribution and habitat
In East Africa, lion populations are present in Somalia, Ethiopia, Kenya, Tanzania, Uganda and Rwanda, and east of the Nile River in Sudan and South Sudan. Due to the civil war in latter two countries, little is known about the conservation status of lions there. In the 1980s, lions used to be present in grasslands and forests of protected areas in both countries. 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.
In Ethiopia, lions are present in Gambella, Omo and Bale Mountains National Parks, around the Chew Bahir and Turkana lakes, and in the Webi Shabeelle area. 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. In 2016, a group of lions was recorded in Alatash National Park close to the international border with Sudan.
In Kenya, lions had been observed near Kavirondo, near Lake Manyara, around Mount Kilimanjaro and in the Tanga Region in the late 19th century. By the 21st century, lion populations in Kenya and Tanzania have been fragmented to 17 patches ranging in size from 86 to 127,515 km2 (33 to 49,234 sq mi). As of 2006, there were an estimated 675 lions in the Tsavo area, 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 3,684 km2 (1,422 sq mi).
In Uganda, lions are present in Kidepo Valley and Murchison Falls National Parks. Those in Queen Elizabeth National Park form a contiguous population with about 60 Central African lions in Virunga National Park in the northeastern part of the Democratic Republic of the Congo. 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.
The lion range in 28 East African protected areas totals 780,400 km2 (301,300 sq mi), of which the following complexes are considered lion strongholds:
- Ruaha National Park cum Rungwa Game Reserve
- Serengeti National Park cum Maasai Mara
- Tsavo East National Park, Tsavo West National Park with Mkomazi National Park
- Selous Game Reserve
The lion population in what used to be the Natal and Cape Provinces of South Africa had been locally extinct since the 1850s to late 1860s. The last lions south of the Orange River were sighted between 1850 and 1858. Eventually, lions were relocated to Addo Elephant National Park.
Elsewhere in Southern Africa, lions are confined to 23 unfenced and 16 fenced reserves; 10 of the fenced reserves are located in South Africa. Lion populations are also present in Namibia, Angola and northern Botswana. In the southwestern part of the Democratic Republic of the Congo, they are considered regionally extinct.
The lion range in nine Southern African protected areas totals 1,540,171 km2 (594,663 sq mi), of which the following protected area complexes are considered lion strongholds:
- North Luangwa National Park and South Luangwa National Park
- Kgalagadi Transfrontier Park
- Okavango Delta cum Hwange National Park
- Great Limpopo Transfrontier Park
- Niassa Reserve
- Zambezi National Park with adjacent protected areas along Zambezi River in Zambia and Mozambique
Lions, which used to live in southern parts of Central Africa, like Gabon and the Republic of the Congo do genetically not fall within the Central African lion but belong to the Southern lion. In Gabon, the presence of lions in Batéké Plateau National Park was doubtful in 2010. In 2015, a camera trap recorded a single male lion in the protected area. Continued camera trapping in the area for more than one year recorded the same lion repeatedly. Hair samples of the lion 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. It is also thought possible that this lion dispersed to the area from Namibia or Botswana.
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.
Behaviour and ecology
The lion is a social cat, living in groups of related individuals with their offspring. Such a family group is called a pride. The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have also been observed. Male lion groups are called a coalition. Membership only changes with the births and deaths of female lions. Male cubs are excluded from their maternal pride when they reach maturity at around 2–3 years of age. The sole known exception of this pattern is the Tsavo lion pride, which always has just one adult male.
Male lions spend years in a nomadic phase before gaining residence in a pride. A study in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.
Lions usually hunt in groups and prey foremost on ungulates such as wildebeest, zebra, African buffalo, gemsbok and giraffe. 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.
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). Predation on adult African bush elephants has been observed in Chobe National Park, Botswana. Sympatric predators include the leopard, cheetah, hyena and African wild dog.
Attacks on humans
- In the 19th century, north of Bechuanaland, a lion non-fatally attacked David Livingstone, who was defending a sheep in a village.
- Two Tsavo males have been known as man-eaters, after an incident during the building of the Uganda Railway in the 1890s. Their skulls and skins are part of the zoological collection of the Field Museum of Natural History in Chicago, the United States of America. The total number of people killed is unclear, but allegedly 135 people fell victim to these lions in less than a year before Colonel John Patterson killed them.
- The "Njombe lions" were a pride of lions in Njombe, in what was then Tanganyika, which for over three generations are thought to have preyed on 1,500 to 2,000 people. They were eventually dispatched by George Rushby.
- In February 2018, a suspected poacher was killed and eaten by lions near Kruger National Park.
- Towards the end of the same month, conservationist Kevin Richardson took three lions for a walk at Dinokeng Game Reserve, near Pretoria in South Africa. A lioness then pursued an impala for at least 2 km (1.2 mi), before unexpectedly killing a 22-year-old woman near her car.
- In July 2018, a "loud commotion" coming from lions was heard by an anti-poaching dog in Sibuya Game Reserve near Kenton-on-Sea, South Africa. The next day, human remains were found in the lion enclosure. They were suspected to have been rhino-poachers, as they had equipment such as a high-powered rifle and wire cutters.
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. Alhough 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.
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.
African lion populations are included in CITES Appendix II. In several South African countries local communities generate significant revenue through wildlife tourism, which is a strong incentive for their support of conservation measures.
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. More than 2000 lions exist in the well-protected Kruger National Park. In June 2015, seven lions were relocated from KwaZulu-Natal in South Africa to Akagera National Park in Rwanda.
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.
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. Interest in the Cape lion had led to attempts to conserve possible descendants in places like Tygerberg Zoo.
Male at Kgalagadi Transfrontier Park, South Africa
Lioness at Phinda Private Game Reserve
Lion pair at Selous Game Reserve, Tanzania
A male in Murchison Falls National Park, Uganda
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