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Lions are divided into two subspecies,[1] which are further divided into genetic clades which overlap in certain parts of Africa to form mixed populations.[2] Panthera leo leo inhabits northern parts of Africa, and Panthera leo melanochaita inhabits southern parts of Africa.[1] Places where genetic analyses demonstrate genetic admixture, or for which results regarding the phylogeography of lions were mixed, are northern parts of East Africa and northern parts of Central Africa.[2][3] Former synonyms include P. l. kamptzi,[4][5][6] P. l. azandica,[4][7] P. l. abyssinica, P. l. massaica, P. l. nubica, P. l. roosevelti, P. l. somaliensis, and P. l. webbiensis.[8]

Northeast / East African population

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Male at Addis Ababa Zoo, Ethiopia

In Northeast Africa, lions occur in Ethiopia, Somalia, South Sudan and Sudan, but are regionally extinct in Djibouti and Eritrea.[9] The results of genetic tests published in 2016 indicated that lions in the region where North Africa overlaps with East Africa (including northern Kenya and possibly northern Uganda), are genetically mixed[2] between the Central African (P. l. leo) and Southern African lions (P. l. melanochaita). As a consequence, the taxonomic status of lions in the Horn of Africa was not resolved by the Cat Specialist Group.[1] Subpopulations were referred to by trinomina such as Panthera leo nubica, Panthera leo roosevelti and Panthera leo somaliensis, and names such as "Abyssinian lion", "Egyptian lion", "Nubian lion", "Somali lion" and "Sudan lion",[10] depending on the locality of occurrence.[8][6][11]

Genetic analyses and taxonomic history

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Young male Nubian lion in New York Zoo, 1903[12]
Abyssinian lion in New York Zoo, 1914[13]

In the 19th century, a number of subspecies were described for lions in Northeast Africa. For example, zoological specimens from Nubia and Somalia were described or proposed by zoologists under the trinomina Felis leo nubicus[14] and Felis leo somaliensis.[15] In later centuries, these trinomina were alternatively considered to be synonymous with the scientific names of the North[16][17] and East African lions.[8][11]

A test done in 2012 on 15 lions at Addis Ababa Zoo and lions from 6 wild populations demonstrated that the captive lions were genetically different to wild lions in other parts of East Africa, but similar to wild lions from Cameroon and Chad.[18][19]

Among six samples from captive lions which were of Ethiopian origin, five samples clustered with other East African samples, but one clustered with Sahelian samples.[20] For a subsequent study, also eight additional samples from wild lions from the Ethiopian Highlands were included in a subsequent analysis. Three of them clustered with the Central African lion clade, and five with other East African samples. The Ethiopian Highlands east and west of the Rift Valley was therefore assumed by scientists to have been a zone of genetic admixture between the two phylogeographic groups.[2]

Results of a phylogeographic analysis using samples from African and Asiatic lions was published in 2006. One of the African samples was a vertebra from the National Museum of Natural History (France) that originated in the Nubian part of Sudan. In terms of mitochondrial DNA, it grouped with lion skull samples from the Central African Republic, Ethiopia and the northern part of the Democratic Republic of the Congo.[3]

A phylogeographic analysis of Pleistocene cave lions, the results of which were published in 2009, revealed that a lion sample from Sudan was distinct from lion samples that originated in the northeastern part of Congo-Kinshasa.[21]

In 2016, analysis of the genomes of lions showed that there had been a basal split between lions in northern and southern parts of Africa. Lions in northern Central Africa belong to the northern clade, and those in Southeast Africa belonged to the southern clade. Lions samples from Ethiopia were shown to be related to both the South-East African and Central African groups, indicating an overlap between these groups there.[2]

In 2017, the Cat Classification Taskforce of the Cat Specialist Group subsumed lion populations in West, Central and North Africa to P. l. leo, and those in East Africa to P. l. melanochaita. At the same time, it was stated that these two subspecies overlap in Ethiopia.[1]

One of the largest lion populations in Ethiopia is found in Gambella. According to genetic research, this population, which is contigous with populations in Sudan, does not belong to the Southern subspecies but to the Northern lion. The same is probably true for the populations in northern Ethiopia,[2] where, a group of lions was recorded in 2016 in Alatash National Park close to the international border with Sudan.[22][23][24]

Other parts of Ethiopia, which still have lions fall into the admixture zone. These are Omo and Bale Mountains National Parks, the ara around the Chew Bahir and Turkana lakes, and the Webi Shabeelle area.[25] 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.[26]

Lions of northern Uganda have not been analysed genetically,[2] and might belong to the Northern subspecies. In northern Uganda, lions are present in Kidepo Valley and Murchison Falls National Parks.[25][27]

In captivity

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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.[19] [28]

Central African population

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A Congolese lioness (formerly P. l. azandica)[4][7] with a partial mane in Virunga National Park, which is adjacent to Queen Elizabeth National Park in Uganda. The Southeast African lion (P. l. melanochaita)[1] is believed to be in parts of D. R. Congo or Central Africa that are adjacent to East African countries like Uganda.[2]
Captive Cameroon lions (formerly P. l. kamptzi)[5][6] at Mefou National Park, Central Africa. Cameroon lions belong to the Central clade of the northern subspecies[2] (P. l. leo).[1]
Further information: Central African lion

The Central African lion[29] is a population of lions in Central Africa that has been grouped under the northern subspecies (Panthera leo leo), but was also found to be related to the southern subspecies[3][2] (Panthera leo melanochaita),[1][9] depending on the subpopulation, and is fragmented into small and isolated groups since the 1950s.[30][25]

In 2010, Gabon, the presence of lions in Gabon's Batéké Plateau National Park had been doubtful.[31] In 2015, a camera trap recorded a single male lion in this national park.[32] Continued camera trapping in the area for more than one year recorded the same lion repeatedly. Its hair samples were collected for phylogenetic analysis by Barnett et al., and compared with tissue samples of lions from Gabon and the Republic of the Congo that were killed in the 20th century. Results indicate that this individual, besides extinct lions in Odzala-Kokoua National Park in the Republic of the Congo, is closely related to the ancestral lion population of the area, and that its DNA shows a typical Southern lion haplotype. It is considered possible that this lion dispersed to the area from Namibia or Botswana.[3]

By contrast, a phylogeographical analysis conducted by Bertola et al. suggested that lions in northern Central Africa, such as Cameroon, belong to the Northern lion group.[2]

In captivity

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In 2006, 1258 captive lions were registered in the International Species Information System, including 13 individuals originating from Senegal to Cameroon, and 970 with uncertain origin.[33]

See also

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References

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  1. ^ a b c d e f g 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.
  2. ^ a b c d e f g h i j k 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. doi:10.1038/srep30807. PMC 4973251. PMID 27488946.
  3. ^ a b c d 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): 1–8. doi:10.1007/s10592-017-1039-2. PMC 6448349. PMID 31007636.
  4. ^ a b c Allen, J. A. (1924). "Carnivora Collected By The American Museum Congo Expedition". Bulletin of the American Museum of Natural History. 47: 73–281.
  5. ^ a b Matschie, P. (1900). "Einige Säugethiere aus dem Hinterlande von Kamerun". Sitzungs-Berichte der Gesellschaft der Naturforschender Freunde zu Berlin. 3: 87–100.
  6. ^ a b c Heptner, V. G.; Sludskij, A. A. (1992) [1972]. "Lion". Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola [Mammals of the Soviet Union. Volume II, Part 2. Carnivora (Hyaenas and Cats)]. Washington DC: Smithsonian Institution and the National Science Foundation. pp. 83–95. ISBN 978-90-04-08876-4.
  7. ^ a b Jackson, D. (2010). "Introduction". Lion. London: Reaktion Books. pp. 1–21. ISBN 978-1861897350.
  8. ^ a b c Haas, S.K.; Hayssen, V.; Krausman, P.R. (2005). "Panthera leo" (PDF). Mammalian Species. 762: 1–11. doi:10.1644/1545-1410(2005)762[0001:PL]2.0.CO;2. S2CID 198968757. Archived from the original (PDF) on 28 July 2017.
  9. ^ a b IUCN Red List of Threatened Species. 2017-3. 2016. 2016. doi:10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en https://www.iucnredlist.org/species/15951/107265605. {{cite journal}}: Missing or empty |title= (help); Unknown parameter |assessor2= ignored (help); Unknown parameter |assessor3= ignored (help); Unknown parameter |assessor4= ignored (help); Unknown parameter |assessor5= ignored (help); Unknown parameter |assessor= ignored (help); Unknown parameter |taxon= ignored (help)
  10. ^ Pease, A. E. (1909-10-16). The Book of the Lion. Ravenio Books.
  11. ^ a b Jonathan Kingdon; David Happold; Thomas Butynski; Michael Hoffmann; Meredith Happold; Jan Kalina (2013-05-23). Mammals of Africa. Vol. 1–6. A & C Black. p. 150. ISBN 978-1-4081-8996-2.
  12. ^ "List of members", Eighth Annual Report, The New York Zoological Society, 1903, p. 32
  13. ^ "Summary of membership", Nineteenth Annual Report, The New York Zoological Society, 1914, p. 36
  14. ^ 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 (in French). Vol. 2. Paris: J. B. Baillière et Fils. p. 186.
  15. ^ Noack, T. (1891). "Felis leo". Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten. 9 (1): 120.
  16. ^ Allen, G. M. (1939). "A Checklist of African Mammals". Bulletin of the Museum of Comparative Zoology at Harvard College. 83: 1–763.
  17. ^ 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.
  18. ^ "A New, Genetically Distinct Lion Population is Found". News Watch. National Geographic Society. 30 November 2012. Retrieved 13 December 2015. The Addis Ababa zoo lions have dark manes and small bodies, unlike other African lions. But life in captivity can sometimes influence appearance. A team of researchers, led by the Max Planck Institute for Evolutionary Anthropology in Germany and the University of York in the UK, checked to see if the lions really are different by comparing DNA samples of 15 lions from the zoo to six populations of wild lions. Their genetic analysis revealed that the gene sequence of all fifteen lions were unique and showed little sign of inbreeding. The study was recently published in the European Journal of Wildlife Research.
  19. ^ a b 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. S2CID 508478.
  20. ^ 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. S2CID 82728679.
  21. ^ Barnett, R.; Shapiro, B.; Barnes, I. A. N.; Ho, S. Y. W.; Burger, J.; Yamaguchi, N.; Higham, T. F. G.; Wheeler, H. T.; Rosendahl, W.; Sher, A. V.; Sotnikova, M.; Kuznetsova, T.; Baryshnikov, G. F.; Martin, L. D.; Harington, C. R.; Burns, J. A.; Cooper, A. (2009). "Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity". Molecular Ecology. 18 (8): 1668–1677. doi:10.1111/j.1365-294X.2009.04134.x. PMID 19302360. S2CID 46716748.
  22. ^ "Lions rediscovered in Ethiopia's Alatash National Park". BBC News. 2016. Retrieved 1 February 2016.
  23. ^ Howard, B. C. (2016). "Once Thought Extinct, 'Lost' Group of Lions Discovered in Africa". National Geographic. Retrieved 2016-02-07.
  24. ^ "Hidden population of up to 200 lions found in remote Ethiopia". New Scientist. 2016. Retrieved 2 February 2016.
  25. ^ a b c 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. S2CID 86796885.
  26. ^ Yirga, G.; Gebresenbet, F.; Deckers, J.; 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. doi:10.4314/mejs.v6i2.109714.
  27. ^ Riggio, J.; Jacobson, A.; Dollar, L.; Bauer, H.; Becker, M.; Dickman, A.; Funston, P.; Groom, R.; Henschel, P.; De Iongh, H.; Lichtenfeld, L.; Pimm, S. (2013). "The size of savannah Africa: a lion's (Panthera leo) view". Biodiversity Conservation. 22 (1): 17–35. doi:10.1007/s10531-012-0381-4. S2CID 18891375.
  28. ^ 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. S2CID 24543070.
  29. ^ Pocock, R. I. (1939). "Panthera leo". The Fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. London: Taylor and Francis Ltd. pp. 212–222.
  30. ^ Chardonnet, P. (2002). Conservation of African lion (PDF). Paris: International Foundation for the Conservation of Wildlife. Archived from the original (PDF) on 10 November 2013.
  31. ^ Henschel, P. H.; Azani, D. E.; Burton, C. O.; Malanda, G.; Saidu, Y. O.; Sam, M. O.; Hunter, L. U. (2010). "Lion status updates from five range countries in West and Central Africa". Cat News. 52: 34–39.
  32. ^ Hedwig, D.; Kienast, I.; Bonnet, M.; Curran, B. K.; Courage, A.; Boesch, C.; Kühl, H. S.; King, T. (2017). "A camera trap assessment of the forest mammal community within the transitional savannah‐forest mosaic of the Batéké Plateau National Park, Gabon". African Journal of Ecology. 56 (4): 777–790. doi:10.1111/aje.12497.
  33. ^ 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. S2CID 24190889. Archived from the original (PDF) on 2006-08-24.


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