Panthera leo melanochaita
|Panthera leo melanochaita|
|Male lion in Etosha National Park, Namibia|
|Female lion in Samburu National Reserve, Kenya|
P. l. melanochaita
|Panthera leo melanochaita|
(Ch. H. Smith, 1842)
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, and 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. 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.
The type specimen 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 and the Republic of the Congo indicate their close genetic relation to P. l. melanochaita samples from Namibia and Botswana.
Charles Hamilton Smith described the type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus. In the 19th and 20th centuries, several naturalists described zoological specimens from Southern and East Africa and proposed the following subspecies:
- Felis leo somaliensis (Noack 1891), based on two lion specimens from Somalia
- Felis leo massaicus (Neumann 1900), based on two lions killed near Kibaya and the Gurui River in Kenya
- Felis leo sabakiensis (Lönnberg 1910), based on two male lions from the environs of Mount Kilimanjaro
- Felis leo roosevelti (Heller 1913), a lion from the Ethiopian Highlands presented to Theodore Roosevelt
- Felis leo nyanzae (Heller 1913), a lion skin from Kampala, Uganda
- Felis leo bleyenberghi (Lönnberg 1914), a male lion from the Katanga Province of Belgian Congo
- Leo leo hollisteri (Joel Asaph Allen 1924), a male lion from the area of Lime Springs, Sotik on the eastern shore of Lake Victoria
- Leo leo krugeri (Austin Roberts 1929), an adult male lion from the Sabi Sand Game Reserve named in honour of Paul Kruger
- 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, 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.
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 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 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, 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. 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.
Since the beginning of the 21st century, several phylogenetic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Scientists analysed between 32 and 480 lion samples from up to 22 countries. 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. It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.
Among six samples from captive lions that originated in Ethiopia, five samples clustered with samples from East Africa, but one clustered with samples from the Sahel. For a subsequent phylogeographic study, also eight wild lion samples from the Ethiopian Highlands were included in the analysis using 194 lion sequences from 22 countries. Four wild lion samples from Ethiopia clustered with lion samples from Central Africa, and four with samples from East Africa, indicating that the Great Rift Valley was not a complete barrier to gene flow. Southeastern Ethiopia is therefore considered a genetic admixture zone between Central and East African lions.
Lions samples from Gabon's Batéké Plateau National Park and Odzala-Kokoua National Park in Republic of the Congo were found to be genetically closely related to lion samples from Namibia and Botswana.
The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. In the 19th and 20th centuries, lion type specimens were described on the basis of mane size and colour. Mane colour varies between tawny, isabelline and light reddish yellow. The Cape lion type specimen had a black mane extending beyond the shoulders and under the belly. Yet, black-maned lions also occur in the Kalahari and eastern Okavango Delta alongside those with a normal tawny colour. 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. 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. In ancient Egyptian art, lions are depicted with weak manes. The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations. Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.
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. 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 an altitude of 800 m (2,600 ft) 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.
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. 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.
Size and weight
Average head-to-body length of male lions is 2.47–2.84 m (97–112 in) with an average weight of 187–193 kg (412–425 lb) in Southern Africa and 174.9 kg (386 lb) in East Africa, while females average 124.2–139.8 kg (274–308 lb) in Southern Africa and 119.5 kg (263 lb) in East Africa. The largest known lion measured 3.30 m (10.8 ft).
In 1936, a man-eating lion shot 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.
Distribution and habitat
In East and Southern Africa, lion populations declined in:
- Somalia since the early 20th century. Intensive poaching since the 1980s and civil unrest posed a threat to lion persistence.
- Uganda 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 in northeastern Democratic Republic of the Congo.
- Kenya 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 Amboseli−Tsavo West−Tsavo 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 3,684 km2 (1,422 sq mi). Lion populations in Kenya and Tanzania are fragmented over 17 patches ranging in size from 86 to 127,515 km2 (33 to 49,234 sq mi).
- Rwanda and Tanzania due to killing of lions during the Rwandan Civil War and ensuing refugee crisis 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 and Zambia due to illegal hunting of prey species in protected areas.
- Botswana 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.
- Namibia 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 Parks.
- 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. 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 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. In Venetia Limpopo Nature Reserve, 18 lions were trophy hunted and 11 euthanized 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.
|Range countries||Lion Conservation Units||Area in km2|
|Democratic Republic of Congo||Massif D'itombwe, Luama||8,441|
|Democratic Republic of Congo, Uganda||Queen Elizabeth−Virunga||5,583|
|Uganda||Toro−Semulik, Lake Mburo, Murchison Falls||4,800|
|Kenya||Laikipia−Samburu, Meru and Nairobi National Parks||43,706|
|Kenya, Tanzania||Serengeti−Mara and Tsavo−Mkomazi||75,068|
|Tanzania||Dar-Biharamulo, Ruaha−Rungwa, Mpanga-Kipengere, Tarangire, Wami Mbiki−Saadani, Selous||384,489|
|Mozambique||Cahora Bassa, Gilé, Gorongosa−Marromeu||82,715|
|Mozambique, Zambia||Middle Zambezi||64,672|
|Mozambique, South Africa||Great Limpopo Transfrontier Park||150,347|
|Zambia||Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex||72,569|
|Zambia, Malawi||North−South Luangwa||72,992|
|Botswana, South Africa||Kgalagadi||163,329|
|Angola||Kissama−Mumbondo, Bocoio−Camacuio, Alto Zambeze||393,760|
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.
Behaviour and ecology
In the Serengeti National Park, 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 2,000 km2 (770 sq mi). Of these, 35 coalitions included male lions that were born in the area but had left and returned after about two years of absence. Nomadic coalitions became resident at between 3.5 and 7.3 years of age.
Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002.
In the Kavango–Zambezi Transfrontier Conservation Area, lions have been monitored since 1999. In 2003, 50 lions were radio-collared in Hwange National Park and tracked until 2012. Results show that adult male and female lions preferred grassland and shrubland habitat, but avoided woodlands and areas with high human density. By contrast, subadult dispersing male lions avoided grasslands and shrublands, but moved in human-dominated areas to a larger extent. Hence, dispersing lions are more vulnerable to coming into conflict with humans than adult lions. In the semi-arid savanna of Zimbabwe’s Hwange National Park, 19 lions were radio-collared and tracked between 2002 and 2007. Both female and male lions moved foremost within 2 km (1.2 mi) of waterholes in all seasons.
Hunting and diet
Lions usually hunt in groups and prey foremost on ungulates such as gemsbok (Oryx gazella), Cape buffalo (Syncerus caffer), blue wildebeest (Connochaetes taurinus), giraffe (Giraffa camelopardalis), common eland (Tragelaphus oryx), greater kudu (T. strepsiceros), nyala (T. angasii), roan antelope (Hippotragus equinus), sable antelope (H. niger), plains zebra (Equus quagga), bushpig (Potamochoerus larvatus), common warthog (Phacochoerus africanus), hartebeest (Alcephalus buselaphus), common tsessebe (Damaliscus lunatus), waterbuck (Kobus ellipsiprymnus), kob (K. kob) and Thomson's gazelle (Eudorcas thomsonii). Their prey is usually in the range of 190–550 kg (420–1,210 pounds). 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).
In Botswana's Chobe National Park, lions also prey on 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, lions killed eight elephants aged between one and 11 years, and two of them older than eight years.
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 in Chicago. The total number of people killed is unclear, but allegedly 135 people fell victim to these lions in less than a year before Colonel Patterson killed them.
- The Njombe lions were a lion pride in Njombe in former Tanganyika, which are thought to have preyed on 1,500 to 2,000 people. They were eventually killed by George Gilman Rushby.
- Between 1990 and 2004, lions killed more than 560 people in Tanzania, mostly during harvest time 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 took three lions for a walk at Dinokeng Game Reserve in South Africa. A lioness pursued an impala for at least 2 km (1.2 mi), and killed a young woman near her car.
- 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.
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. 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.
African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes. 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. 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.
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, 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. In addition, about 100 captive lions were registered in ISIS as P. l. krugeri, which derived from lions captured in South Africa. Interest in the Cape lion had led to attempts to conserve possible descendants in Tygerberg Zoo.
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 the 'southern subspecies'.
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- Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O’Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z.; Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News. Special Issue 11: 71−73.
- Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P.; Nowell, K. (2016). "Panthera leo". The IUCN Red List of Threatened Species. IUCN. 2016: e.T15951A115130419. doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en.
- Bauer, H.; Chapron, G.; Nowell, K.; Henschel, P.; Funston, P.; Hunter, L. T.; Macdonald, D. W.; Packer, C. (2015). "Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas". Proceedings of the National Academy of Sciences. 112 (48): 14894–14899. doi:10.1073/pnas.1500664112. PMC 4672814. PMID 26504235.
- IUCN Cat Specialist Group (2006). Conservation Strategy for the Lion Panthera leo in Eastern and Southern Africa (PDF). Pretoria, South Africa: IUCN.
- Mazak, V. (1975). "Notes on the Black-maned Lion of the Cape, Panthera leo melanochaita (Ch. H. Smith, 1842) and a Revised List of the Preserved Specimens". Verhandelingen Koninklijke Nederlandse Akademie van Wetenschappen (64): 1–44.
- Barnett, R.; Sinding, M. H.; Vieira, F. G.; Mendoza, M. L.; Bonnet, M.; Araldi, A.; Kienast, I.; Zambarda, A.; Yamaguchi, N.; Henschel, P.; Gilbert, M. T. (2018). "No longer locally extinct? Tracing the origins of a lion (Panthera leo) living in Gabon". Conservation Genetics. 19 (3): 611–618. doi:10.1007/s10592-017-1039-2.
- Smith, C. H. (1842). "Black maned lion Leo melanochaitus". In Jardine, W. (ed.). The Naturalist's Library. Vol. 15 Mammalia. London: Chatto and Windus. p. Plate X, 177.
- Noack, T. (1891). "Felis leo". Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten. 9 (1): 120.
- Neumann, O. (1900). "Die von mir in den Jahren 1892–95 in Ost- und Central-Afrika, speciell in den Massai-Ländern und den Ländern am Victoria Nyansa gesammelten und beobachteten Säugethiere". Zoologische Jahrbücher. Abtheilung für Systematik, Geographie und Biologie der Thiere. 13 (VI): 529–562.
- Lönnberg, E. (1910). "Mammals". In Sjöstedt, Y. (ed.). Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906. Volume 1. Stockholm: P. Palmquists Aktiebolag. pp. 1−58, Plate 1−7.
- Heller, E. (1913). "New races of carnivores and baboons from equatorial Africa and Abyssinia". Smithsonian Miscellaneous Collections. 61 (19): 1–12.
- Lönnberg, E. (1914). "New and rare mammals from Congo". Revue de Zoologie Africaine (3): 273–278.
- Allen, J. A. (1924). "Carnivora Collected By The American Museum Congo Expedition". Bulletin of the American Museum of Natural History. 47: 73–281.
- Roberts, A. (1929). "New forms of African mammals". Annals of the Transvaal Museum. 21 (13): 82–121.
- Roberts, A. (1948). "Descriptions of some new subspecies of mammals". Annals of the Transvaal Museum. 21 (1): 63–69.
- Zukowsky, L. (1964). "Eine neue Löwenrasse als weiterer Beleg für die Verzwergung der Wirbeltierfauna des afrikanischen Osthorns". Milu, Wissenschaftliche und Kulturelle Mitteilungen aus dem Tierpark Berlin (1): 269–273.
- Allen, G. M. (1939). A Checklist of African Mammals. Bulletin of the Museum of Comparative Zoology at Harvard College. 83. Cambridge, Mass. : The Museum. pp. 1–763.
- Lundholm, B. (1952). "A skull of a Cape lioness (Felis leo melanochaitus H. Smith". Annals of the Transvaal Museum (32): 21–24.
- Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung. 17: 167–280.
- Wozencraft, W.C. (2005). "Panthera leo". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 546. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Pocock, R. I. (1930). "The lions of Asia". Journal of the Bombay Natural Historical Society. 34: 638–665.
- Ellerman, J. R.; Morrison-Scott, T. C. S. (1966). "Subgenus Leo Oken, 1816, (Brehm, 1829)". Checklist of Palaearctic and Indian Mammals 1758 to 1946 (Second ed.). London: British Museum (Natural History). p. 319.
- Meester, J.; Setzer, H. W. (1977). The mammals of Africa. An identification manual. Washington, DC: Smithsonian Institution Press.
- Yamaguchi, N. (2000). "The Barbary lion and the Cape lion: their phylogenetic places and conservation" (PDF). African Lion Working Group News. 1: 9–11.
- Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation" (PDF). Conservation Genetics. 7 (4): 507–514. doi:10.1007/s10592-005-9062-0. Archived from the original (PDF) on 2006-08-24.
- O’Brien, S. J.; Martenson, J. S.; Packer, C.; Herbst, L.; de Vos, V.; Joslin, P.; Ott-Joslin, J.; Wildt, D. E. & Bush, M. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions" (PDF). National Geographic Research. 3 (1): 114–124. Archived from the original (PDF) on 2013-05-02.
- Antunes, A.; Troyer, J. L.; Roelke, M. E.; Pecon-Slattery, J.; Packer, C.; Winterbach, C.; Winterbach, H.; Johnson, W. E. (2008). "The Evolutionary Dynamics of the Lion Panthera leo Revealed by Host and Viral Population Genomics". PLOS Genetics. 4 (11): e1000251. doi:10.1371/journal.pgen.1000251. PMC 2572142. PMID 18989457.
- Mazák, J. H. (2010). "Geographical variation and phylogenetics of modern lions based on craniometric data". Journal of Zoology. 281 (3): 194−209.
- Bertola, L. D.; Van Hooft, W. F.; Vrieling, K.; Uit De Weerd, D. R.; York, D. S.; Bauer, H.; Prins, H. H. T.; Funston, P. J.; Udo De Haes, H. A.; Leirs, H.; Van Haeringen, W. A.; Sogbohossou, E.; Tumenta, P. N.; De Iongh, H. H. (2011). "Genetic diversity, evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa" (PDF). Journal of Biogeography. 38 (7): 1356–1367. doi:10.1111/j.1365-2699.2011.02500.x.
- Dubach, J. M., Briggs, M. B., White, P. A., Ament, B. A., & Patterson, B. D. (2013). "Genetic perspectives on "Lion Conservation Units" in Eastern and Southern Africa". Conservation Genetics. 14 (4): 741−755. doi:10.1007/s10592-013-0453-3.CS1 maint: Uses authors parameter (link)
- Bertola, L. D., Tensen, L., Van Hooft, P., White, P. A., Driscoll, C. A., Henschel, P., Caragiulo, A., Dias-Freedman, I., Sogbohossou, E. A., Tumenta, P. N. and Jirmo, T. H. (2015). "Autosomal and mtDNA markers affirm the distinctiveness of lions in West and Central Africa". PLOS One. 10 (10): e0137975. Bibcode:2015PLoSO..1037975B. doi:10.1371/journal.pone.0137975. PMC 4605676. PMID 26466139.CS1 maint: Uses authors parameter (link)
- Bertola, L. D.; Jongbloed, H.; Van Der Gaag, K. J.; De Knijff, P.; Yamaguchi, N.; Hooghiemstra, H.; Bauer, H.; Henschel, P.; White, P. A.; Driscoll, C. A. & Tende, T. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports. 6: 30807. Bibcode:2016NatSR...630807B. doi:10.1038/srep30807. PMC 4973251. PMID 27488946.
- Guggisberg, C. A. W. (1975). "Lion Panthera leo (Linnaeus, 1758)". Wild Cats of the World. New York: Taplinger Publishing. pp. 138–179. ISBN 978-0-8008-8324-9.
- Guggisberg, C. A. W. (1963). Simba: The Life of the Lion. Philadelphia, PA: Chilton Books. ASIN B000OKBJQ0.
- Smithers, R. H. N. (1971). "Panthera leo". The Mammals of Botswana. Pretoria: University of Pretoria. pp. 116−119.
- Nagel, D.; Hilsberg, S.; Benesch, A.; Scholtz, J. (2003). "Functional morphology and fur patterns in recent and fossil Panthera species". Scripta Geologica. 126: 227–239.
- Bruche, S.; Gusset, M.; Lippold, S.; Barnett, R.; Eulenberger, K.; Junhold, J.; Driscoll, C. A.; Hofreiter, M. (2012). "A genetically distinct lion (Panthera leo) population from Ethiopia". European Journal of Wildlife Research. 59 (2): 215–225. doi:10.1007/s10344-012-0668-5.
- West, P. M.; Packer, C. (2002). "Sexual Selection, Temperature, and the Lion's Mane". Science. 297 (5585): 1339–1943. Bibcode:2002Sci...297.1339W. doi:10.1126/science.1073257. PMID 12193785.
- Gnoske T. P.; Celesia, G. G.; Kerbis, Peterhans J. C. (2006). "Dissociation between mane development and sexual maturity in lions (Panthera leo): solution to the Tsavo riddle?". Journal of Zoology. 270 (4): 551–560. Bibcode:2010JZoo..281..263G. doi:10.1111/j.1469-7998.2006.00200.x.
- Turner, J. A.; Vasicek, C. A.; Somers, M. J. (2015). "Effects of a colour variant on hunting ability: the white lion in South Africa". Open Science Repository Biology: e45011830.
- McBride, C. (1977). The White Lions of Timbavati. Johannesburg: E. Stanton. ISBN 978-0-949997-32-6.
- Tucker, L. (2003). Mystery of the White Lions—Children of the Sun God. Mapumalanga: Npenvu Press. ISBN 978-0-620-31409-1.
- Smuts, G. L.; Robinson, G. A.; Whyte, I. J. (1980). "Comparative growth of wild male and female lions (Panthera leo)". Journal of Zoology. 190 (3): 365–373. Bibcode:2010JZoo..281..263G. doi:10.1111/j.1469-7998.1980.tb01433.x.
- Nowell, K.; Jackson, P. (1996). "African lion" (PDF). Wild Cats: Status Survey and Conservation Action Plan. Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21. ISBN 978-2-8317-0045-8.
- Wood, G. L. (1983). The Guinness Book of Animal Facts and Feats. Sterling Publishing. ISBN 978-0-85112-235-9.
- Riggio, J.; Jacobson, A.; Dollar, L.; Bauer, H.; Becker, M.; Dickman, A.; Funston, P.; Groom, R.; Henschel, P.; De Iongh, H.; Lichtenfeld, L. (2013). "The size of savannah Africa: a lion's (Panthera leo) view". Biodiversity and Conservation. 22 (1): 17–35. doi:10.1007/s10531-012-0381-4.
- Funaioli, U. and Simonetta, A. M. (1966). "The Mammalian Fauna of the Somali Republic: Status and Conservation Problems". Monitore Zoologico Italiano, Supplemento. 1 (1): 285–347. doi:10.1080/03749444.1966.10736746.CS1 maint: Multiple names: authors list (link)
- Chardonnet, P. (2002). "Chapter II: Population Survey" (PDF). Conservation of the African Lion : Contribution to a Status Survey. Paris: International Foundation for the Conservation of Wildlife, France & Conservation Force, USA. pp. 21–101. Archived from the original (PDF) on 10 November 2013.
- Fagotto, F. (1985). "Larger animals of Somalia in 1984". Environmental Conservation. 12 (3): 260−264. doi:10.1017/S0376892900016015.
- Treves, A.; Naughton-Treves, L. (1999). "Risk and opportunity for humans coexisting with large carnivores". Journal of Human Evolution. 36 (3): 275−282. doi:10.1006/jhev.1998.0268. PMID 10074384.
- Omoya, E. O.; Mudumba, T.; Buckland, S. T.; Mulondo, P. & Plumptre, A. J. (2014). "Estimating population sizes of lions Panthera leo and spotted hyaenas Crocuta crocuta in Uganda's savannah parks, using lure count methods" (PDF). Oryx. 48 (3): 394–401. doi:10.1017/S0030605313000112.
- Treves, A.; Plumptre, A. J.; Hunter, L. T. B.; Ziwa, J. (2009). "Identifying a potential lion Panthera leo stronghold in Queen Elizabeth National Park, Uganda, and Parc National des Virunga, Democratic Republic of Congo". Oryx. 43: 60−66. doi:10.1017/S003060530700124X.
- Frank, L., Maclennan, S., Hazzah, L., Hill, T., Bonham, R. (2006). Lion Killing in the Amboseli-Tsavo Ecosystem, 2001–2006, and its Implications for Kenya's Lion Population (PDF). Nairobi: Living with Lions.CS1 maint: Uses authors parameter (link)
- Dolrenry, S., Hazzah, L. and Frank, L. G. (2016). "Conservation and monitoring of a persecuted African lion population by Maasai warriors". Conservation Biology. 30 (3): 467−475. doi:10.1111/cobi.12703. PMID 27111059.CS1 maint: Multiple names: authors list (link)
- Dolrenry, S., Stenglein, J., Hazzah, L., Lutz, R. S. and Frank, L. (2014). "A metapopulation approach to African lion (Panthera leo) conservation". PLOS One. 9 (2): e88081. Bibcode:2014PLoSO...988081D. doi:10.1371/journal.pone.0088081. PMC 3914926. PMID 24505385.CS1 maint: Multiple names: authors list (link)
- Bauer, H.; Van Der Merwe, S. (2004). "Inventory of free-ranging lions Panthera leo in Africa". Oryx. 38 (1): 26–31. doi:10.1017/S0030605304000055.
- Bantlin, D. A. (2017). Movements of Reintroduced Lions in Akagera National Park, Rwanda in Relation to Prey Abundance (Doctoral dissertation). Madison: University of Wisconsin−Madison.
- Snyman, A., Jackson, C.R. and Funston, P. J. (2015). "The effect of alternative forms of hunting on the social organization of two small populations of lions Panthera leo in Southern Africa". Oryx. 49 (4): 604−610. doi:10.1017/S0030605313001336.CS1 maint: Multiple names: authors list (link)
- Stander, P. E. (1990). "A suggested management strategy for stock-raiding lions in Namibia". South African Journal of Wildlife Research. 20 (2): 37−43.
- Ferreira, S. M.; Govender, D.; Herbst, M. (2013). "Conservation implications of Kalahari lion population dynamics". African Journal of Ecology. 51 (2): 176–179. doi:10.1111/aje.12003.
- Rautenbach, I. L. (1978). "Panthera leo (Linnaeus, 1758)". The Mammals of the Transvaal. Pietermaritzburg: University of Natal. pp. 471−475.
- Hayward, M. W., Adendorff, J., O’Brien, J., Sholto-Douglas, A., Bissett, C., Moolman, L. C., Bean, P., Fogarty, A., Howarth, D., Slater, R. and Kerley, G. I. (2007). "Practical considerations for the reintroduction of large, terrestrial, mammalian predators based on reintroductions to South Africa's Eastern Cape Province" (PDF). The Open Conservation Biology Journal. 1 (1): 1−11. doi:10.2174/1874839200701010001.CS1 maint: Uses authors parameter (link)
- Mudumba, T.; Mulondo, P.; Okot, E.; Nsubuga, M.; Plumptre, A. (2009). National Census of Lions and Hyaenas in Uganda. Strategic Action Plan for Large Carnivore Conservation in Uganda. Kampala: Uganda Wildlife Authority.
- Mésochina, P.; Mbangwa, O.; Chardonnet, P.; Mosha, R.; Mtui, B.; Drouet, N.; Kissui, B. (2010). Conservation status of the lion (Panthera leo Linnaeus, 1758) in Tanzania. Paris: SCI Foundation, MNRT-WD, TAWISA & IGF Foundation.
- Chardonnet, P.; Mésochina, P.; Bento, C.; Conjo, D.; Begg, C. l. (2009). Conservation status of the lion (Panthera leo Linnaeus, 1758) in Mozambique (PDF). Maputo, Mozambique: Fondation IGF, Campfire Association, SCI Foundation, The Ratel Trust, Direcção Nacional de Terras e Florestas.
- Schaller, G. B. (1972). The Serengeti lion: A study of predator–prey relations. Chicago: University of Chicago Press. ISBN 978-0-226-73639-6.
- Bertram, B.C.R. (1973). "Lion population regulation". East Africa Wildlife Journal. 11 (3–4): 215−225. doi:10.1111/j.1365-2028.1973.tb00088.x.
- Bertram, B. (1975). "Social factors influencing reproduction in wild lions". Journal of Zoology. 177 (4): 463–482. doi:10.1111/j.1469-7998.1975.tb02246.x.CS1 maint: Uses authors parameter (link)
- Bygott, J. D., Bertram, B. C. R., and Hanby, J. P. (1979). "Male lions in large coalitions gain reproductive advantages". Nature. 282 (282): 839–841. Bibcode:1979Natur.282..839B. doi:10.1038/282839a0.CS1 maint: Uses authors parameter (link)
- Borrego, N., Ozgul, A., Slotow, R. and Packer, C. (2018). "Lion population dynamics: do nomadic males matter?". Behavioral Ecology. 29 (3): 660–666. doi:10.1093/beheco/ary018.CS1 maint: Multiple names: authors list (link)
- Patterson, B. D., Kasiki, S. M., Selempo, E. and Kays, R. W. (2004). "Livestock predation by lions (Panthera leo) and other carnivores on ranches neighboring Tsavo National Parks, Kenya". Biological Conservation. 119 (4): 507−516. CiteSeerX 10.1.1.884.8117. doi:10.1016/j.biocon.2004.01.013.CS1 maint: Multiple names: authors list (link)
- Elliot, N. B.; Cushman, S. A.; Macdonald, D. W.; Loveridge, A. J. (2014). "The devil is in the dispersers: predictions of landscape connectivity change with demography". Journal of Applied Ecology. 51 (5): 1169–1178. doi:10.1111/1365-2664.12282.
- Valeix, M., Loveridge, A. J., Davidson, Z., Madzikanda, H., Fritz, H. and Macdonald, D. W. (2010). "How key habitat features influence large terrestrial carnivore movements: waterholes and African lions in a semi-arid savanna of north-western Zimbabwe". Landscape Ecology. 25 (3): 337–351. doi:10.1007/s10980-009-9425-x.CS1 maint: Multiple names: authors list (link)
- Hayward, M. W.; Kerley, G. I. H. (2005). "Prey preferences of the lion (Panthera leo)" (PDF). Journal of Zoology. 267 (3): 309–322. Bibcode:2010JZoo..281..263G. CiteSeerX 10.1.1.611.8271. doi:10.1017/S0952836905007508.
- Davidson, Z., Valeix, M., Van Kesteren, F., Loveridge, A.J., Hunt, J.E., Murindagomo, F. and Macdonald, D.W. (2013). "Seasonal diet and prey preference of the African lion in a waterhole-driven semi-arid savanna". PLOS One. 8 (2): e55182. doi:10.1371/journal.pone.0055182.g001. PMC 3566210. PMID 23405121.CS1 maint: Multiple names: authors list (link)
- Joubert, D. (2006). "Hunting behaviour of lions (Panthera leo) on elephants (Loxodonta africana) in the Chobe National Park, Botswana". African Journal of Ecology. 44 (44): 279–281. doi:10.1111/j.1365-2028.2006.00626.x.
- Power, R. J.; Compion, R. X. S. (2009). "Lion predation on elephants in the Savuti, Chobe National Park, Botswana". African Zoology. 44 (1): 36–44. doi:10.3377/004.044.0104.
- Kerbis Peterhans, J. C. and Gnoske, T. P. (2001). "The Science of 'Man-Eating*' Among Lions Panthera leo With a Reconstruction of the Natural History of the 'Man-Eaters of Tsavo'". Journal of East African Natural History. 90 (1): 1–40. doi:10.2982/0012-8317(2001)90[1:TSOMAL]2.0.CO;2.CS1 maint: Multiple names: authors list (link)
- Patterson, J. H. (1907). The man-eaters of Tsavo and other East African adventures. London: Macmillan and Co.
- Rushby, G. G. (1965). No More the Tusker. London: W. H. Allen.
- Packer, C., Ikanda, D., Kissui, B. and Kushnir, H. (2005). "Lion attacks on humans in Tanzania" (PDF). Nature. 436 (7052): 927−928. Bibcode:2005Natur.436..791S. doi:10.1038/436791a. PMID 16094358.CS1 maint: Uses authors parameter (link)
- "South African lions eat 'poacher', leaving just his head". The BBC. 2018. Retrieved 2018-02-25.
- Haden, A. (2018). "Suspected poacher mauled to death by lions close to Kruger National Park". The South African. Retrieved 25 February 2018.
- Torchia, C. (2018). "Lion kills woman at refuge of South African 'lion whisperer'". The Washington Post. Retrieved 28 February 2018.
- "Lions eat 'rhino poachers' on South African game reserve". BBC. 2018. Retrieved 2018-07-19.
- Williams, V. L.; Loveridge, A. J.; Newton, D. J.; Macdonald, D. W. (2017). "A roaring trade? The legal trade in Panthera leo bones from Africa to East-Southeast Asia". PLOS One. 12 (10): e0185996. Bibcode:2017PLoSO..1285996W. doi:10.1371/journal.pone.0185996. PMC 5655489. PMID 29065143.
- Schroeder, R. A. (2018). "Moving Targets: The 'Canned'Hunting of Captive-Bred Lions in South Africa". African Studies Review. 61 (1): 8–32. doi:10.1017/asr.2017.94.
- "'Herdsmen' poison lions, vultures — but not in Nigeria". The Cable. 2018-02-17. Retrieved 2018-02-24.
- Kamoga, J. (2018). "East African lions dying of poisoning". The Observer. Retrieved 2018-02-24.
- Winter, S. (2018-02-16). "Lion MASSACRE as six big cats die after eating 'poison'". Daily Express. Retrieved 2018-02-24.
- "Zimbabwe's 'iconic' lion Cecil killed by hunter". BBC News. 27 July 2015. Archived from the original on 28 July 2015. Retrieved 3 August 2015.
- "Xanda, son of Cecil the lion, killed by hunter in Zimbabwe". BBC News. 20 July 2017. Retrieved 20 July 2017.
- Midlane, N. (2013). The conservation status and dynamics of a protected African lion Panthera leo population in Kafue National Park, Zambia. Cape Town: University of Cape Town.
- Tefera, M. (2003). "Phenotypic and reproductive characteristics of lions (Panthera leo) at Addis Ababa Zoo". Biodiversity & Conservation. 12 (8): 1629–1639. doi:10.1023/A:1023641629538.
- "South Africa: Lion Cubs Thought to Be Cape Lions". Associated Press Archive. 2000.
- "'Extinct' lions (Cape lion) surface in Siberia". The BBC News. 2000. Retrieved 2012-12-31.
- Jackson, D. (2010). "Introduction". Lion. London: Reaktion Books. pp. 1–21. ISBN 978-1861897350.
- Borzo, G. (2002). "Unique social system found in famous Tsavo lions". EurekAlert.
- Hunter, L.; Barrett, P. (2018). "Lion Panthera leo". The Field Guide to Carnivores of the World (Second ed.). London, Oxford, New York, New Delhi, Sydney: Bloomsbury. pp. 46−47. ISBN 978-1-4729-5080-2.
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