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Lions in [[Queen Elizabeth National Park]] form a contiguous population with lions in [[Virunga National Park]] in the northeastern part of the [[Democratic Republic of the Congo]],<ref name=Riggioetal2013/><ref name="Treves_al2009"/><ref name="Bauer_vanderMerwe">{{Cite journal |last1=Bauer |first1=H. |last2=Van Der Merwe |first2=S. |doi=10.1017/S0030605304000055 |title=Inventory of free-ranging lions ''Panthera leo'' in Africa |journal=Oryx |volume=38 |year=2004 |issue=1 |pages=26–31}}</ref> 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.<ref name="Omoyaetal2014">{{cite journal |author=Omoya, E. O. |author2=Mudumba, T. |author3=Buckland, S. T. |author4=Mulondo, P. |author5=Plumptre, A. J. |last-author-amp=yes |year=2014 |title=Estimating population sizes of lions ''Panthera leo'' and spotted hyaenas ''Crocuta crocuta'' in Uganda's savannah parks, using lure count methods |journal=Oryx |volume=48 |issue=3 |pages=394–401 |url=https://research-repository.st-andrews.ac.uk/bitstream/handle/10023/6885/omoya2014oryx394.pdf?sequence=1}}</ref>
Lions in [[Queen Elizabeth National Park]] form a contiguous population with lions in [[Virunga National Park]] in the northeastern part of the [[Democratic Republic of the Congo]],<ref name=Riggioetal2013/><ref name="Treves_al2009"/><ref name="Bauer_vanderMerwe">{{Cite journal |last1=Bauer |first1=H. |last2=Van Der Merwe |first2=S. |doi=10.1017/S0030605304000055 |title=Inventory of free-ranging lions ''Panthera leo'' in Africa |journal=Oryx |volume=38 |year=2004 |issue=1 |pages=26–31}}</ref> 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.<ref name="Omoyaetal2014">{{cite journal |author=Omoya, E. O. |author2=Mudumba, T. |author3=Buckland, S. T. |author4=Mulondo, P. |author5=Plumptre, A. J. |last-author-amp=yes |year=2014 |title=Estimating population sizes of lions ''Panthera leo'' and spotted hyaenas ''Crocuta crocuta'' in Uganda's savannah parks, using lure count methods |journal=Oryx |volume=48 |issue=3 |pages=394–401 |url=https://research-repository.st-andrews.ac.uk/bitstream/handle/10023/6885/omoya2014oryx394.pdf?sequence=1}}</ref>

As of 2006, there were an estimated 675 lions in the [[Tsavo]] area, out of the 2,000 total in Kenya.<ref name="Frank_al2006">{{cite book |author1=Frank, L. |author2=Maclennan, S. |author3=Hazzah, L. |author4=Hill, T. |author5=Bonham, R. |year=2006 |title=Lion Killing in the Amboseli-Tsavo Ecosystem, 2001–2006, and its Implications for Kenya's Lion Population |url=http://www.livingwithlions.org/AnnualReports/2006-Lion-killing-in-Amboseli-Tsavo-ecosystem.pdf |publisher=Living with Lions |location=[[Nairobi]], Kenya}}</ref> Between 2004 and 2013, lion guardians around [[Amboseli National Park]] identified 65 lions in an area of {{convert|3684|km2|sqmi|abbr=on}}.<ref>{{cite journal |author=Dolrenry, S., Hazzah, L. and Frank, L. G. |year=2016 |title=Conservation and monitoring of a persecuted African lion population by Maasai warriors |volume=30 |journal=Conservation Biology |issue=3 |pages=467−475 |doi= 10.1111/cobi.12703}}</ref>
As of 2006, there were an estimated 675 lions in the [[Tsavo]] area, out of the 2,000 total in Kenya.<ref name="Frank_al2006">{{cite book |author1=Frank, L. |author2=Maclennan, S. |author3=Hazzah, L. |author4=Hill, T. |author5=Bonham, R. |year=2006 |title=Lion Killing in the Amboseli-Tsavo Ecosystem, 2001–2006, and its Implications for Kenya's Lion Population |url=http://www.livingwithlions.org/AnnualReports/2006-Lion-killing-in-Amboseli-Tsavo-ecosystem.pdf |publisher=Living with Lions |location=[[Nairobi]], Kenya}}</ref> Between 2004 and 2013, lion guardians around [[Amboseli National Park]] identified 65 lions in an area of {{convert|3684|km2|sqmi|abbr=on}}.<ref>{{cite journal |author=Dolrenry, S., Hazzah, L. and Frank, L. G. |year=2016 |title=Conservation and monitoring of a persecuted African lion population by Maasai warriors |volume=30 |journal=Conservation Biology |issue=3 |pages=467−475 |doi= 10.1111/cobi.12703}}</ref> Lion populations in Kenya and Tanzania are fragmented over 17 patches ranging in size from {{convert|86|to|127515|km2|sqmi|abbr=on}}.<ref>{{cite journal |author=Dolrenry, S., Stenglein, J., Hazzah, L., Lutz, R. S. and Frank, L. |year=2014 |title=A metapopulation approach to African lion (''Panthera leo'') conservation |journal= PLOS ONE |volume=9 |issue=2 |pages=e88081 |url=http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0088081|doi=10.1371/journal.pone.0088081 |pmid=24505385 |pmc=3914926 |bibcode=2014PLoSO...988081D}}</ref>
A small population is present in Rwanda's [[Akagera National Park]], estimated at 35 individuals at most in 2004.<ref name="Bauer_vanderMerwe"/>
A small population is present in Rwanda's [[Akagera National Park]], estimated at 35 individuals at most in 2004.<ref name="Bauer_vanderMerwe"/>

Revision as of 17:44, 2 December 2018

Panthera leo melanochaita
Male lion in Etosha National Park, Namibia
Female lion in Samburu National Reserve, Kenya
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. leo
Subspecies:
P. l. melanochaita
Trinomial name
Panthera leo melanochaita
(Ch. H. Smith, 1842)
Synonyms[1]

formerly:

  • P. l. bleyenberghi
  • P. l. krugeri
  • P. l. vernayi
  • P. l. massaica
  • P. l. nyanzae

Panthera leo melanochaita is a lion subspecies in Southern and East Africa.[2] 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.[3] 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.[4] In 2005, a Lion Conservation Strategy was developed for East and Southern Africa.[5]

The type specimen for P. l. melanochaita was a black-maned lion from the Cape of Good Hope, known as the Cape lion.[6]

Taxonomic history

Lions shot in Kenya's Sotik Plains in 1909

Charles Hamilton Smith described the type specimen for Panthera leo melanochaita in 1842 using the scientific name Felis (Leo) melanochaitus.[7] In the 19th and 20th centuries, several naturalists described zoological specimens from Southern and East Africa and proposed the following subspecies:

Dispute over the validity of these purported subspecies continued among naturalists and curators of natural history museums until the early 21st century.[6][17][18][19][1]

In the 20th century, some authors supported the view of the Cape lion being a distinct subspecies.[14][17][18][20] 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.[17]

Pocock subordinated lions to the genus Panthera in 1930, when he wrote about Asiatic lions.[21] 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.[22] Various authors recognized between seven and 10 African lion subspecies.[19] Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.[23]

In the 1970s, the scientific name P. l. vernayi was considered synonymous with P. l. krugeri.[19] In 1975, Vratislav Mazák hypothesized that the Cape lion evolved geographically isolated from other populations by the Great Escarpment.[6] 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.[24][25]

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.[1] In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo.[3] 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.[2]

Genetic research

Range map including proposed clades and the two subspecies (P. l. leo and P. l. melanochaita) according to genetic research

Since the beginning of the 21st century, several phylogenetic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Scientists analysed between 32 and 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.[26] It was assumed that tropical rainforest and the East African Rift constituted major barriers between the two groups.[25][27][28][29][30][31] Based on this assessment, the species comprises two recognised subspecies:[2]

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

The two groups were in contact in Ethiopia. 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 lions in North, West and Central Africa. In nine of 19 lion samples from Ethiopia, haplotypes of the Central African lion group were found, indicating that the Great Rift Valley was not a complete barrier to gene flow; southeastern Ethiopia is considered a genetic admixture zone between Central and East African lions.[31]

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.[32] 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.[27][29] 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.[27]

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.[33]

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.[34]

Characteristics

Adult male lions in Serengeti National Park, Tanzania
Male lion in Narok County, southern Kenya
A male lion in Amboseli National Park, Kenya
File:White-lion.jpg
White lion

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. Average head-to-body length of male lions is 2.47–2.84 m (97–112 in) with a weight of 148.2–190.9 kg (327–421 lb). The largest East African lion measured 3.33 m (10.9 ft). Females are smaller and less heavy.[35]

An exceptionally heavy male lion near Mount Kenya weighed 272 kg (600 lb).[36] Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa.[37] The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.[33] 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.[38]

Manes

In the 19th and 20th centuries, lion type specimen were described on the basis of mane size and colour. Mane colour varies between tawny, isabelline and light reddish yellow.[39] The Cape lion type specimen had a black mane extending beyond the shoulders and under the belly.[7] Yet, black-maned lions also occur in the Kalahari and eastern Okavango Delta alongside those with a normal tawny colour.[40] 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.[9] 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.[12] A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well.[11] Until the late 20th century, mane colour and size was thought to be a distinct subspecific characteristic.[19][6]

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.[41] 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.[42] Weak manes is also a characteristic feature of lions depicted in ancient Egyptian art.[43]

White lion

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.[44] White lions are selected for breeding in captivity.[45] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during canned hunts.[46]

Distribution and habitat

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

In East and Southern Africa, lion population declined in:

Contemporary lion distribution and habitat quality in East and Southern Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[5] Between 2002 and 2012, educated guesses for size of populations in these LCUs ranged from 33,967 to 32,000 individuals.[49][56]

Range countries Lion Conservation Units Area in km2
Democratic Republic of Congo Massif D'itombwe, Luama 8,441[5]
Democratic Republic of Congo, Uganda Queen Elizabeth-Virunga 5,583[57]
Uganda Toro-Semulik, Lake Mburo, Murchison Falls 4,800[58]
Somalia Arboweerow-Alafuuto 24,527[5]
Somalia, Kenya Bushbush-Arawale 22,540[5]
Kenya Laikipia-Samburu, Meru and Nairobi National Parks 43,706[56]
Kenya, Tanzania Serengeti-Mara and Tsavo-Mkomazi 75,068[47]
Tanzania Dar-Biharamulo, Ruaha-Rungwa, Mpanga-Kipengere, Tarangire, Wami Mbiki-Saadani, Selous 384,489[47]
Tanzania, Mozambique Niassa 177,559[59]
Mozambique Cahora Bassa, Gilé, Gorongosa-Marromeu 82,715[59]
Mozambique, Zambia Middle Zambezi 64,672[59]
Mozambique, South Africa Great Limpopo Transfrontier Park 150,347[59]
Zambia Liuwa Plains, Sioma Ngwezi, Kafue Sumbu Complex 72,569[56]
Zambia, Malawi North-South Luangwa 72,992[56]
Malawi Kasungu, Nkhotakota 4,187[56]
Zimbabwe Mapungubwe, Bubye 10,033[56]
Botswana, Zimbabwe Okavango-Hwange 99,552[56]
Botswana Xaixai 12,484[5]
Botswana, South Africa Kgalagadi 163,329[56]
Angola Kissama-Mumbondo, Bocoio-Camacuio, Alto Zambeze 393,760[5]
Angola, Namibia Etosha-Kunene 123,800[5]
Namibia Khaudum-Caprivi 92,372[5]

The LCUs Ruaha-Rungwa, Serengeti-Mara, Tsavo-Mkomazi and Selous in East Africa, as well as Luangwa, Kgalagadi, Okavango-Hwange, Mid-Zambezi, Niassa and Greater Limpopo in Southern Africa are currently considered as lion strongholds. These LCUs host more than 500 individuals each, and the population trend is stable there.[56]

Lions in Queen Elizabeth National Park form a contiguous population with lions in Virunga National Park in the northeastern part of the Democratic Republic of the Congo,[56][57][60] 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.[61]

As of 2006, there were an estimated 675 lions in the Tsavo area, out of the 2,000 total in Kenya.[62] Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of 3,684 km2 (1,422 sq mi).[63] 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).[64]

A small population is present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004.[60]

The lion population in South Africa's former Natal and Cape Provinces is locally extinct since the mid 19th century.[65] The last lions south of the Orange River were sighted between 1850 and 1858.[6] Between 2000 and 2004, 34 lions were reintroduced to eight protected areas in the Eastern Cape Province, including Addo Elephant National Park.[66]

Behaviour and ecology

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

In the Serengeti National Park, monitoring of lion prides started in 1966.[67] Between 1966 and 1972, two observed lion prides comprised between seven and 10 females each. Females had litters once in 23 months on average.[68] 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.[69] Male lions in coalitions are closely related.[70] 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.[71]

Lions living near ranches in the vicinity of Tsavo East National Park consisted of three prides, two pairs and a single lion in 2002.[72]

Results of a 10-year long survey on 50 radio-collared lions in the Kavango–Zambezi Transfrontier Conservation Area show that adult lions preferred grassland and shrubland habitat, but avoided woodlands and areas with high human density; by contrast, subadult dispersing male lions avoided grasslands and shrublands, but moved in human-dominated areas to a larger extent. Hence, dispersing lions are more vulnerable to coming into conflict with humans than adult lions.[73] 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.[74]

Hunting and diet

Lions usually hunt in groups and prey foremost on ungulates such as 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 to 270 kg (88 to 595 pounds).[75] 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.[67]

In Botswana's Chobe National Park, lions also prey on African bush elephants .[76][77]

Attacks on humans

Threats

In Africa, lions are threatened by pre-emptive killing or in retaliation for preying on livestock. Prey base depletion, loss and conversion of habitat have led to a number of subpopulations becoming small and isolated. Trophy hunting has contributed to population declines in Botswana, Namibia, Zimbabwe and Zambia.[3] 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.[62]

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.[88] Private game ranches in South Africa also breed lions for the canned hunting industry.[89]

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle.[90] 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.[91][92]

In 2015 and 2017, two male lions, Cecil and his son Xanda, were killed by trophy hunters in Zimbabwe's Hwange National Park.[93][94]

In Zambia's Kafue National Park, uncontrolled bushfires and hunting of lions and prey species makes it difficult for the lion population to recover. Cub mortality in particular is high.[95]

Conservation

African lions are included in CITES Appendix II. Today, lion populations are stable only in large protected area complexes.[3] 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.[5] 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.[3]

In captivity

Captive lion in Philadelphia Zoo

At the beginning of the 21st century, the Addis Ababa Zoo kept 16 adult lions. It is assumed that their ancestors, five males and two females, were caught in southwestern Ethiopia as part of a zoological collection for Emperor Haile Selassie I.[33][96]

In 2006, the International Species Information System (ISIS) showed 29 lions that had derived from lions captured in Angola and Zimbabwe. In addition, about 100 captive lions were registered in ISIS as P. l. krugeri, which derived from lions captured in South Africa.[25] Interest in the Cape lion had led to attempts to conserve possible descendants in places like Tygerberg Zoo.[97][98]

Regional names

Lion populations in Southern and East Africa were referred to by several regional names, including "Katanga lion", "Transvaal lion", "Kalahari lion",[11][14][15] "Southeast African lion", and "Southwest African lion",[99] "Masai lion", "Serengeti lion,"[67] "Tsavo lion"[100] and "Uganda lion".[19]

See also

References

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  2. ^ a b c 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.
  3. ^ a b c d e Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P.; Nowell, K. (2016). "Panthera leo". The IUCN Red List of Threatened Species. 2016. IUCN: e.T15951A115130419. doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en.{{cite iucn}}: error: |doi= / |url= mismatch (help)
  4. ^ 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.
  5. ^ a b c d e f g h i j IUCN Cat Specialist Group (2006). Conservation Strategy for the Lion Panthera leo in Eastern and Southern Africa (PDF). Pretoria, South Africa: IUCN.
  6. ^ a b c d e f 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.
  7. ^ a b 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.
  8. ^ Noack, T. (1891). "Felis leo". Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten. 9 (1): 120.
  9. ^ a b 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.
  10. ^ 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. {{cite book}}: External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help)
  11. ^ a b c Lönnberg, E. (1914). "New and rare mammals from Congo". Revue de Zoologie Africaine (3): 273–278.
  12. ^ a b c Heller, E. (1914). "New races of carnivores and baboons from equatorial Africa and Abyssinia". Smithsonian Miscellaneous Collections. 61 (19): 1–12.
  13. ^ Allen, J. A. (1924). "Carnivora Collected By The American Museum Congo Expedition". Bulletin of the American Museum of Natural History. 47: 73–281.
  14. ^ a b c Roberts, A. (1929). "New forms of African mammals". Annals of the Transvaal Museum. 21 (13): 82–121.
  15. ^ a b Roberts, A. (1948). "Descriptions of some new subspecies of mammals". Annals of the Transvaal Museum. 21 (1): 63–69.
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