East African lion

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East African lion
Lions @ Maasai Mara (20792033896).jpg
Male at Maasai Mara, Serengeti part of Kenya
Lioness, the Serengeti (13) (28007348834).jpg
Female (lioness) in Serengeti National Park, Tanzania
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species: P. leo
Subspecies: P. l. melanochaita[1]
Trinomial name
Panthera leo melanochaita[1]
(Ch. H. Smith, 1842)
Synonyms[2]

formerly:

  • P. l. massaica
  • P. l. hollisteri
  • P. l. nyanzae

The East African lion (Panthera leo melanochaita)[1] occurs in East Africa, but is regionally extinct in Djibouti and Eritrea.[3][4] Lion populations in the East African range countries have increasingly declined since the turn of the century.[5]

Formerly, the East African lion was divided into several subspecies.[2][6] Results of a phylogeographic analysis indicate that lion populations in East African and Southern Africa range countries are genetically close, forming a distinct clade.[7] In 2017, lion populations in this part of Africa were subsumed to P. l. melanochaita.[1]

Taxonomic history[edit]

In the 19th and 20th centuries, several lion type specimens from East Africa were described and proposed as subspecies:

In 1939, the American zoologist Allen recognized the trinomina F. l. hollisteri, F. l. nyanzae and F. l. massaica as valid subspecies in East Africa. He considered F. l. nubicus and F. l. somaliensis as synonymous with F. l. leo, but F. l. sabakiensis and F. l. roosevelti as synonyms of F. l. massaica.[16]

Pocock subordinated lions to the genus Panthera in 1930, when he wrote about Asiatic lions.[17] 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.[18] Various authors recognized between seven and 10 African lion subspecies.[15] Others followed the classification proposed by Ellerman and Morrison-Scott, recognizing two subspecies including one in Africa.[19]

In 2005, P. l. massaica, P. l. hollisteri and P. l. nyanzae were considered valid taxa.[2] In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo.[4] In 2017, the number of valid lion subspecies was reduced to two, namely P. l. leo in India, North, West and Central Africa, and P. l. melanochaita in East and Southern Africa.[1]

Genetic research[edit]

Results of genetic analysis indicate that East and Southern African lions form a clade distinct from the North African lion.[20]

Since 2005, several phylogeographic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Five lion samples from Kenya's Tsavo East National Park showed identical haplotypes as three lion samples from the Transvaal region in South Africa.[21] 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.[22] 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.[23]

Characteristics[edit]

Adult male lions with long brown manes in Serengeti National Park
Male with a partially black mane in Narok County, southern Kenya
A mature male lion with an intermediate mane development in Amboseli National Park, Kenya

The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. Average head-to-body length of male lions is 2.47–2.84 m (8.1–9.3 ft) with a weight of 148.2–190.9 kg (327–421 lb). The largest East African lion measured 3.33 m (10.9 ft). Females are smaller and less heavy.[24] An exceptionally heavy male near Mount Kenya weighed 272 kg (600 lb).[25]

Male lions killed in East Africa were less heavy than lions killed by hunters in Southern Africa.[26] The captive male lions at Addis Ababa Zoo have darker manes and smaller bodies than those of wild populations.[22]

Manes[edit]

In the 19th and 20th centuries, lion type specimen were described on the basis of mane size and colour.[27] 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.[28] 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.[29]

A male lion specimen from Somalia had a short mane.[15] 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.[13] A few lions observed in the environs of Mount Kilimanjaro had tawny to sandy coloured manes as well.[12] 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.[11] Mane colour of males in Kenya vary between tawny, isabelline and light reddish yellow.[27] 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.[30] 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.[31]

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.[32]

Distribution and habitat[edit]

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.[3]

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.[33]

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.[34] 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.[35] In 2016, a group of lions was recorded in Alatash National Park close to the international border with Sudan.[36][37][38]

In Kenya, lions had been observed near Kavirondo, near Lake Manyara, around Mount Kilimanjaro and in the Tanga Region in the late 19th century.[11] 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).[39] As of 2006, there were an estimated 675 lions in the Tsavo area, out of the 2,000 total in Kenya.[40] Between 2004 and 2013, lion guardians around Amboseli National Park identified 65 lions in an area of 3,684 km2 (1,422 sq mi).[41]

In Uganda, lions are present in Kidepo Valley and Murchison Falls National Parks.[34][42] 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.[42][43][34] 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.[44]

A small population is present in Rwanda's Akagera National Park, estimated at 35 individuals at most in 2004.[34]

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:[42]

Ecology and behaviour[edit]

Mating at the Ngorongoro Conservation Area, Tanzania
Pride in Serengeti National Park
At a Cape buffalo kill in the Serengeti

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.[45] The sole known exception of this pattern is the Tsavo lion pride, which always has just one adult male.[46]

Male lions spend years in a nomadic phase before gaining residence in a pride.[47] A study in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[48]

Lions usually hunt in groups and prey foremost on ungulates such as wildebeest, zebra, African buffalo, gemsbok and giraffe.[49] 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.[45][47]

Man-eating[edit]

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.[50][51] 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.[52]

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.[53]

Threats[edit]

In Africa, local people kill lions pre-emptively or in retaliation for preying on livestock. Lion populations are also threatened by depletion of prey base, loss and conversion of habitat.[4] Though lions and their prey are officially protected in Tsavo, they are regularly killed by local people, with over 100 known lion killings between 2001 and 2006.[40]

In 2014, seven lions in Ikona Wildlife Management Area were reportedly poisoned by a herdsman for attacking his cattle.[54] 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.[55][56]

Conservation[edit]

African lion populations are included in CITES Appendix II. In several East African countries local communities generate significant revenue through wildlife tourism, which is a strong incentive for their support of conservation measures.[4]

In captivity[edit]

A male at Addis Ababa Zoo, Ethiopia, 2006

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.[57]

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.[58][22]

Gallery[edit]

Cultural significance[edit]

The Prudhoe Lions from around 1370 BC

The lion is featured as an animal symbol in East Africa.[59][60] The name Simba is a Swahili word for the lion, which also means 'aggressive', 'king' and 'strong'.[61]

See also[edit]

References[edit]

  1. ^ a b c d 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). 
  2. ^ a b c Wozencraft, W.C. (2005). "Panthera leo". In Wilson, D.E.; Reeder, D.M. 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. 
  3. ^ a b Chardonnet, P. (2002). "Chapter II: Population Survey". Conservation of the African Lion : Contribution to a Status Survey (PDF). Paris: International Foundation for the Conservation of Wildlife, France & Conservation Force, USA. pp. 21–101. Archived from the original (PDF) on 10 November 2013. 
  4. ^ a b c d Bauer, H.; Packer, C.; Funston, P. F.; Henschel, P.; Nowell, K. (2016). "Panthera leo". IUCN Red List of Threatened Species. Version 2017-3. International Union for Conservation of Nature.  doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en 10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en
  5. ^ Bauer, H., Chapron, G., Nowell, K., Henschel, P., Funston, P., Hunter, L.T., Macdonald, D.W. and 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. Bibcode:2015PNAS..11214894B. doi:10.1073/pnas.1500664112. PMC 4672814Freely accessible. 
  6. ^ Stott, K. (1950). "Locality records of African mammals". Journal of Mammalogy. 31 (2): 185–189. doi:10.2307/1375515. 
  7. ^ 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.; Ottosson, U.; Saidu, Y.; Vrieling, K.; de Iongh, H. H. (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. 
  8. ^ Blainville, H. M. D. de (1843). "F. leo nubicus". Ostéographie ou description iconographique comparée du squelette et du système dentaire des mammifères récents et fossils pour servir de base à la zoologie et la géologie. Vol 2. Paris: J. B. Baillière et Fils. p. 186. 
  9. ^ Noack, T. (1891). "Felis leo". Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten. 9 (1): 120. 
  10. ^ a b Allen, J. A. (1924). "Carnivora Collected By The American Museum Congo Expedition". Bulletin of the American Museum of Natural History. 47: 73–281. 
  11. ^ a b c 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. 
  12. ^ a b Lönnberg, E. (1910). "Mammals". In Sjöstedt, Y. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906. Volume 1. Uppsala: Königlich Schwedische Akademie der Wissenschaften. 
  13. ^ a b Heller, E. (1914). "New races of carnivores and baboons from equatorial Africa and Abyssinia". Smithsonian Miscellaneous Collections. 61 (19): 1–12. 
  14. ^ 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. 
  15. ^ a b c 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. 
  16. ^ Allen, G. M. (1939). A Checklist of African Mammals. Bulletin of the Museum of Comparative Zoology at Harvard College 83: 1–763.
  17. ^ Pocock, R. I. (1930). "The lions of Asia". Journal of the Bombay Natural Historical Society. 34: 638–665. 
  18. ^ Ellerman, J. R. and T. C. S. Morrison-Scott (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. 
  19. ^ Meester, J.; Setzer, H. W. (1977). The mammals of Africa. An identification manual. Washington, DC: Smithsonian Institution Press. 
  20. ^ 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. 
  21. ^ Dubach, J.; Patterson, B. D.; Briggs, M. B.; Venzke, K.; Flamand, J.; Stander, P.; Scheepers, L.; Kays, R. W. (2005). "Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo" (PDF). Conservation Genetics. 6 (1): 15–24. doi:10.1007/s10592-004-7729-6. 
  22. ^ a b c 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. 
  23. ^ 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.; Ottosson, U.; Saidu, Y.; Vrieling, K.; de Iongh, H. H. (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. 
  24. ^ Guggisberg, C. A. W. (1975). "Lion Panthera leo (Linnaeus, 1758)". Wild Cats of the World. New York: Taplinger Publishing. pp. 138–179. ISBN 0-8008-8324-1. 
  25. ^ Nowell, K., Jackson, P. (1996). "African lion". Wild Cats: Status Survey and Conservation Action Plan (PDF). Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21. ISBN 2-8317-0045-0. 
  26. ^ 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. doi:10.1111/j.1469-7998.1980.tb01433.x. 
  27. ^ a b Guggisberg, Charles Albert Walter (1963). Simba: The Life of the Lion. Philadelphia, PA: Chilton Books. ASIN B000OKBJQ0. 
  28. ^ 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. doi:10.1111/j.1469-7998.2006.00200.x. 
  29. ^ West, P. M.; Packer, C. (2002). "Sexual Selection, Temperature, and the Lion's Mane". Science. 297 (5585): 1339–1343. Bibcode:2002Sci...297.1339W. doi:10.1126/science.1073257. ISSN 0036-8075. PMID 12193785. 
  30. ^ Call the Hair Club for Lions. The Field Museum.
  31. ^ Borzo, G. (2002). "Unique social system found in famous Tsavo lions". EurekAlert. 
  32. ^ 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. 
  33. ^ Fagotto F. (1985). "Larger animals of Somalia in 1984". Environmental Conservation 12 (3): 260–264. 
  34. ^ a b c d 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. 
  35. ^ Yirga, G., Gebresenbet, F., Deckers, J. and Bauer, H. (2014). "Status of Lion (Panthera leo) and Spotted Hyena (Crocuta crocuta) in Nechisar National Park, Ethiopia". Momona Ethiopian Journal of Science 6(2): 127−137. 
  36. ^ "Lions rediscovered in Ethiopia's Alatash National Park". BBC News. 2016. Retrieved 1 February 2016. 
  37. ^ Howard, B. C. (2016). "Once Thought Extinct, 'Lost' Group of Lions Discovered in Africa". National Geographic. Retrieved 2016-02-07. 
  38. ^ Wong, S. (2016). "Hidden population of up to 200 lions found in remote Ethiopia". New Scientist. Retrieved 2 February 2016. 
  39. ^ 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 3914926Freely accessible. PMID 24505385. 
  40. ^ a b 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 Living with Lions, Nairobi, Kenya, 9.
  41. ^ Dolrenry S., Hazzah L., 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. 
  42. ^ a b c 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. 
  43. ^ Treves A., Plumptre A. J., Hunter L. T., 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 (01): 60–66. doi:10.1017/s003060530700124x. 
  44. ^ Omoya, E.O., Mudumba, T., Buckland, S.T., Mulondo, P. and 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. 
  45. ^ a b Schaller, G. B. (1972). The Serengeti lion: A study of predator–prey relations. Chicago: University of Chicago Press. ISBN 0-226-73639-3. 
  46. ^ Milius, S. (2002). "Biology: Maneless lions live one guy per pride". Society for Science & the Public. 161 (16): 253. doi:10.1002/scin.5591611614. 
  47. ^ a b Hanby, J. P., and Bygott, J. D. (1979). "Population changes in lions and other predators". In Sinclair, A. R. E.; Norton-Griffiths, M. Serengeti: dynamics of an ecosystem. Chicago: The University of Chicago Press. pp. 249−262. 
  48. ^ Borrego, N., Ozgul, A., Slotow, R. and Packer, C. (2018). "Lion population dynamics: do nomadic males matter?". Behavioral Ecology: early view. doi:10.1093/beheco/ary018. 
  49. ^ Hayward, M. W.; Kerley, G. I. H. (2005). "Prey preferences of the lion (Panthera leo)" (PDF). Journal of Zoology. 267 (3): 309–322. doi:10.1017/S0952836905007508. 
  50. ^ 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. 90 (1): 1–40. doi:10.2982/0012-8317(2001)90[1:TSOMAL]2.0.CO;2. 
  51. ^ "Field Museum uncovers evidence behind man-eating; revises legend of its infamous man-eating lions" (Press release). The Field Museum. 2003. 
  52. ^ Patterson, J.H. (1907). The man-eaters of Tsavo and other East African adventures. London: Macmillan and Co.
  53. ^ Rushby, George G. (1965). No More the Tusker. London: W. H. Allen. 
  54. ^ "'Herdsmen' poison lions, vultures — but not in Nigeria". The Cable. 2018-02-17. Retrieved 2018-02-24. 
  55. ^ Kamoga, J. (2018). "East African lions dying of poisoning". The Observer. Retrieved 2018-02-24. 
  56. ^ Winter, S. (2018-02-16). "Lion MASSACRE as six big cats die after eating 'poison'". The Express. Retrieved 2018-02-24. 
  57. ^ Barnett R., Yamaguchi N., Barnes I., Cooper A. (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)". Proceedings of the Royal Society B. 273 (1598): 2119–2125. doi:10.1098/rspb.2006.3555. PMC 1635511Freely accessible. PMID 16901830. 
  58. ^ Tefera, M. (2003). "Phenotypic and reproductive characteristics of lions (Panthera leo) at Addis Ababa Zoo". Biodiversity & Conservation. 12 (8): 1629–1639. Retrieved 2018-05-13. 
  59. ^ Hogarth, C.; Butler, N. (2004). "Animal Symbolism (Africa)". In Walter, M. N. Shamanism: An Encyclopedia of World Beliefs, Practices, and Culture, Volume 1. pp. 3–6. ISBN 1-57607-645-8. 
  60. ^ Lynch, P. A. (2004). African Mythology A to Z. Infobase Publishing. p. 63. ISBN 0-8160-4892-4. 
  61. ^ Brakefield, T. (1993). "Sociable Simba". Big Cats: Kingdom of Might. Voyageur Press. p. 51−68. ISBN 978-0-89658-329-0. 

Further reading[edit]

External links[edit]

Media related to Panthera leo at Wikimedia Commons