Editing Lion (section)

==Taxonomy==
[[File:Two cladograms for Panthera.svg|thumb|right|The upper cladogram is based on the 2006 study,<ref name=Johnson2006/><ref name="werdelin2009">{{cite journal |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J.  |title=Phylogeny and evolution of cats (Felidae) |journal=Biology and Conservation of Wild Felids |year=2010 |pages=59–82 |url=https://www.researchgate.net/publication/266755142 |access-date=10 February 2019 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}</ref> the lower one on the 2010<ref name=davis2010/> and 2011<ref name=mazak2011/> studies.]]

''Felis leo'' was the [[scientific name]] used by [[Carl Linnaeus]] in 1758, who described the lion in his work ''[[Systema Naturae]]''.<ref name="Linn1758">{{cite book |last=Linnaeus|first= C. |year=1758 |title=Caroli Linnæi Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |volume=Tomus I |edition=decima, reformata |location=Holmiae |publisher=Laurentius Salvius |page=41 |chapter=''Felis leo'' |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}} {{in lang|la}}</ref> The genus name ''Panthera'' was coined by [[Lorenz Oken]] in 1816.<ref>{{cite book |last1=Oken |first1=L. |year=1816 |title=Lehrbuch der Zoologie. 2. Abtheilung |location=Jena |publisher=August Schmid & Comp. |page=1052 |chapter=1. Art, ''Panthera'' |chapter-url=https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052}}</ref> Between the mid-18th and mid-20th centuries, 26 lion [[Zoological specimen|specimen]]s were described and proposed as subspecies, of which 11 were recognised as [[Valid name (zoology)|valid]] in 2005.<ref name=MSW3/> They were distinguished mostly by the size and colour of their manes and skins.<ref name=Hemmer/>

===Subspecies===
In the 19th and 20th centuries, several lion [[type specimen]]s were described and proposed as [[subspecies]], with about a dozen recognised as [[Valid name (zoology)|valid]] [[Taxon|taxa]] until 2017.<ref name=MSW3/> Between 2008 and 2016, [[IUCN Red List]] assessors used only two subspecific names: ''P. l. leo'' for African lion populations, and ''P. l. persica'' for the Asiatic lion population.<ref name=IUCN/><ref name=Breitenmoser2008>{{cite iucn |author=Breitenmoser, U. |author2=Mallon, D. P. |author3=Ahmad Khan, J. |author4=Driscoll, C.  |date=2008 |page=e.T15952A5327221 |title=''Panthera leo'' ssp. ''persica'' |doi=10.2305/IUCN.UK.2008.RLTS.T15952A5327221.en}}</ref><ref name=Henschel2015>{{cite iucn |author=Henschel, P. |author2=Bauer, H. |author3=Sogbohoussou, E. |author4=Nowell, K.  |date=2015 |doi=10.2305/IUCN.UK.2015-2.RLTS.T68933833A54067639.en |title=''Panthera leo'' West Africa subpopulation}}</ref> In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion [[Taxonomy (biology)|taxonomy]], and recognises two subspecies based on results of several [[phylogeographic]] studies on lion [[evolution]], namely:<ref name=catsg>{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O'Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S.  |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |volume=Special Issue 11 |pages=71–73 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=71 |access-date=6 August 2019 |archive-date=17 January 2020 |archive-url=https://web.archive.org/web/20200117172708/https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=71 |url-status=live}}</ref>

*[[Panthera leo leo|''P. l. leo'']] {{small|(Linnaeus, 1758)}} − the [[Nominate subspecies|nominate]] lion subspecies includes the [[Asiatic lion]], the [[Locally extinct|regionally extinct]] [[Barbary lion]], and lion populations in West and northern parts of Central Africa.<ref name=catsg/> [[Synonym (taxonomy)|Synonyms]] include ''P. l. persica'' {{small|(Meyer, 1826)}}, ''P. l. senegalensis'' {{small|(Meyer, 1826)}}, ''P. l. kamptzi'' {{small|([[Paul Matschie|Matschie]], 1900)}}, and ''P. l. azandica'' {{small|([[Joel Asaph Allen|Allen]], 1924)}}.<ref name=MSW3/> It has been called 'northern lion' and 'northern subspecies'.<ref name=Wood1865>{{cite book |last=Wood |first=J. G. |author-link=John George Wood |title=The Illustrated Natural History |volume=((Mammalia, Volume 1)) |publisher=[[Routledge]] |chapter=Felidæ; or the Cat Tribe |pages=129−148 |location=London |chapter-url=https://books.google.com/books?id=v1DPAAAAMAAJ&pg=PA147 |year=1865 |access-date=23 December 2018 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505150653/https://books.google.com/books?id=v1DPAAAAMAAJ&pg=PA147#v=onepage&q&f=false |url-status=live}}</ref><ref name=Hunter2018>{{cite book |last1=Hunter |first1=L. |last2=Barrett |first2=P.  |title=The Field Guide to Carnivores of the World |edition=Second |publisher=Bloomsbury |location=London, Oxford, New York, New Delhi, Sydney |isbn=978-1-4729-5080-2 |date=2018 |chapter=Lion ''Panthera leo'' |chapter-url=https://books.google.com/books?id=4HpxDwAAQBAJ&pg=PA46 |pages=46−47 |access-date=1 December 2018 |archive-date=5 March 2024 |archive-url=https://web.archive.org/web/20240305201610/https://books.google.com/books?id=4HpxDwAAQBAJ&pg=PA46 |url-status=live}}</ref>
*[[Panthera leo melanochaita|''P. l. melanochaita'']] {{small|([[Charles Hamilton Smith|Smith]], 1842)}} − includes the extinct [[Cape lion]] and lion populations in East and Southern African regions.<ref name=catsg/> Synonyms include ''P. l. somaliensis'' {{small|(Noack 1891)}}, ''P. l. massaica'' {{small|([[Oscar Neumann|Neumann]], 1900)}}, ''P. l. sabakiensis'' {{small|([[Einar Lönnberg|Lönnberg]], 1910)}}, ''P. l. bleyenberghi'' {{small|(Lönnberg, 1914)}}, ''P. l. roosevelti'' {{small|([[Edmund Heller|Heller]], 1914)}}, ''P. l. nyanzae'' {{small|(Heller, 1914)}}, ''P. l. hollisteri'' {{small|([[Joel Asaph Allen|Allen]], 1924)}}, ''P. l. krugeri'' {{small|([[Austin Roberts (zoologist)|Roberts]], 1929)}}, ''P. l. vernayi'' {{small|(Roberts, 1948)}}, and ''P. l. webbiensis'' {{small|(Zukowsky, 1964)}}.<ref name=MSW3 /><ref name=Hemmer/> It has been referred to as 'southern subspecies' and 'southern lion'.<ref name=Hunter2018/>

However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates that Central African lions are derived from both northern and southern lions, as they cluster with ''P. leo leo'' in mtDNA-based phylogenies whereas their genomic DNA indicates a closer relationship with P. ''leo melanochaita''.<ref>{{cite journal |last1=de Manuel |first1=M. |last2=Barnett |first2=R.|last3=Sandoval-Velasco |first3=M. |last4=Yamaguchi |first4=N. |last5=Vieira |first5=F. G. |last6=Mendoza |first6=M. L. Z. |last7=Liu |first7=S. |last8=Martin |first8=M. D. |last9=Sinding |first9=M-S. S. |last10=Mak |first10=S. S. T. |last11=Carøe |first11=C. |last12=Liu |first12=S. |last13=Guo |first13=C. |last14=Zheng |first14=J. |last15=Zazula |first15=G. |last16=Baryshnikov |first16=G. |last17=Eizirik |first17=E. |last18=Koepfli |first18=K.-P. |last19=Johnson |first19=W. E. |last20=Antunes |first20=A. |last21=Sicheritz-Ponten |first21=T. |last22=Gopalakrishnan |first22=S. |last23=Larson |first23=G. |last24=Yang |first24=H. |last25=O'Brien |first25=S. J. |last26=Hansen |first26=A. J. |last27=Zhang |first27=G. |last28=Marques-Bonet |first28=T. |last29=Gilbert |first29=M. T. P. |date=2020 |title=The evolutionary history of extinct and living lions |journal= Proceedings of the National Academy of Sciences |volume=117 |issue=20 |pages=10927–10934 |doi=10.1073/pnas.1919423117 |pmid= 32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref>

Lion samples from some parts of the [[Ethiopian Highlands]] cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers, therefore, assume Ethiopia is a contact zone between the two subspecies.<ref name=Bertola2016>{{cite journal |author1=Bertola, L. D. |author2=Jongbloed, H. |author3=Van Der Gaag, K. J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P. A. |author10=Driscoll, C. A. |author11=Tende, T. |author12=Ottosson, U. |author13=Saidu, Y. |author14=Vrieling, K. |author15=de Iongh, H. H. |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807 |pmid=27488946 |pmc=4973251  |bibcode=2016NatSR...630807B}}</ref> [[Genome]]-wide data of a wild-born historical lion sample from Sudan showed that it clustered with ''P. l. leo'' in mtDNA-based phylogenies, but with a high affinity to ''P. l. melanochaita''; this suggested that the taxonomic position of lions in Central Africa may require revision.<ref name=DeManuel_al2020>{{cite journal |author1=de Manuel, M. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T.  |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P. |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |doi=10.1073/pnas.1919423117 |year=2020 |pmid=32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref>

===Fossil records===
Other lion subspecies or [[sister species]] to the modern lion existed in prehistoric times:<ref name="Christiansen08CaveLions">{{cite journal |last1=Christiansen |first1=P. |year=2008 |title=Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters |journal=[[Cladistics (journal)|Cladistics]] |volume=24 |issue=6 |pages=977–992 |doi=10.1111/j.1096-0031.2008.00226.x|pmid=34892880 |s2cid=84497516 |doi-access=free }}</ref>
*''[[Panthera leo sinhaleyus|P. l. sinhaleyus]]'' was a [[fossil]] [[carnassial]] excavated in [[Sri Lanka]], which was attributed to a lion. It is thought to have become extinct around 39,000 years ago.<ref>{{Cite journal|first1=K. |last1=Manamendra-Arachchi |first2=R. |last2=Pethiyagoda |first3=R. |last3=Dissanayake |first4=M. |last4=Meegaskumbura  |year=2005 |title=A second extinct big cat from the late Quaternary of Sri Lanka |journal=[[The Raffles Bulletin of Zoology]] |issue=Supplement 12 |pages=423–434 |url=http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |archive-url=https://web.archive.org/web/20070807215533/http://rmbr.nus.edu.sg/rbz/biblio/s12/s12rbz423-434.pdf |url-status=dead |archive-date=7 August 2007 }}</ref>
*[[Panthera fossilis|''P. fossilis'']] was larger than the modern lion and lived in the [[Middle Pleistocene]]. Bone fragments were excavated in caves in the United Kingdom, Germany, Italy and Czech Republic.<ref name=Marciszak2010>{{cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K.  |year=2010 |title=Two forms of cave lion: Middle Pleistocene ''Panthera spelaea fossilis'' Reichenau, 1906 and Upper Pleistocene ''Panthera spelaea spelaea'' Goldfuss, 1810 from the Bisnik Cave, Poland |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117 |bibcode=2010NJGPA.258..339M |url=https://www.researchgate.net/publication/233669138 |access-date=14 March 2019 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 |url-status=live }}</ref><ref name=Sabol2014>{{cite journal |last=Sabol |first=M. |year=2014 |title=''Panthera fossilis'' (Reichenau, 1906) (Felidae, Carnivora) from Za Hájovnou Cave (Moravia, The Czech Republic): A Fossil Record from 1987–2007 |journal=Acta Musei Nationalis Pragae, Series B, Historia Naturalis |volume=70 |issue=1–2 |pages=59–70 | doi=10.14446/AMNP.2014.59 |doi-access=free }}</ref>
*''[[Panthera spelaea|P. spelaea]]'', or the '''cave lion''', lived in [[Eurasia]] and [[Beringia]] during the [[Late Pleistocene]]. It became extinct due to [[climate warming]] or [[Early human migrations|human expansion]] latest by 11,900 years ago.<ref name=Stuart2011>{{cite journal |last1=Stuart |first1=A. J. |last2=Lister |first2=A. M.  |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–2340 |doi=10.1016/j.quascirev.2010.04.023 |bibcode=2011QSRv...30.2329S}}</ref> Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska.<ref name="Hemmer2011">{{cite journal |author=Hemmer, H. |year=2011 |title=The story of the cave lion – ''Panthera Leo Spelaea'' (Goldfuss, 1810) – A review |journal=Quaternaire |volume=4|pages=201–208|url=https://www.researchgate.net/publication/285886884}}</ref> It likely derived from ''P. fossilis'',<ref name=Barnett2016>{{cite journal |last1=Barnett |first1=R. |last2=Mendoza |first2=M. L. Z. |last3=Soares |first3=A. E. R. |last4=Ho |first4=S. Y. W. |last5=Zazula |first5=G. |last6=Yamaguchi |first6=N. |last7=Shapiro |first7=B. |last8=Kirillova |first8=I. V. |last9=Larson |first9=G. |last10=Gilbert |first10=M. T. P.  |year=2016 |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), resolve its position within the ''Panthera'' cats |journal=Open Quaternary |volume=2 |page=4 |doi=10.5334/oq.24 |url=https://ora.ox.ac.uk/objects/uuid:9d4f84e6-64c6-49fd-a1dc-a981ba7e8028/download_file?file_format=pdf&safe_filename=Larson%2Bet%2Bal%252C%2BMitogenomics%2Bof%2Bthe%2BExtinct%2BCave%2BLion%252C%2BPanthera%2Bspelaea%2B%2528Goldfuss%252C%2B1810%2529%252C%2Bresolve%2Bits%2Bposition%2Bwit.pdf&type_of_work=Journal+article |doi-access=free |hdl=10576/22920 |hdl-access=free |access-date=18 May 2019 |archive-date=3 October 2020 |archive-url=https://web.archive.org/web/20201003085724/https://ora.ox.ac.uk/objects/uuid:9d4f84e6-64c6-49fd-a1dc-a981ba7e8028/download_file?file_format=pdf&safe_filename=Larson%2Bet%2Bal%252C%2BMitogenomics%2Bof%2Bthe%2BExtinct%2BCave%2BLion%252C%2BPanthera%2Bspelaea%2B%2528Goldfuss%252C%2B1810%2529%252C%2Bresolve%2Bits%2Bposition%2Bwit.pdf&type_of_work=Journal+article |url-status=live }}</ref> and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia.<ref name="BurgerJ-Molecular-phylogeny">{{Cite journal |last1=Burger |first1=J. |year=2004 |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |pmid=15012963 |volume=30 |issue=3 |pages=841–849 |doi=10.1016/j.ympev.2003.07.020 |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%202004.pdf |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last2=Rosendahl |first2=W. |last10=Alt |first10=K. W. |bibcode=2004MolPE..30..841B  |url-status=dead |archive-url=https://web.archive.org/web/20070925204424/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%202004.pdf |archive-date=25 September 2007 }}</ref><ref name=Barnett2016/> It is depicted in [[Paleolithic]] cave paintings, ivory carvings, and clay busts.<ref name="Packer00">{{Cite journal |author=Packer, C. |author2=Clottes, J.  |title=When Lions Ruled France |journal=Natural History |volume=109 |issue=9 |pages=52–57 |date=2000 |url=http://www.cbs.umn.edu/sites/cbs.umn.edu/files/public/downloads/When_lions_ruled_France.pdf |archive-url=https://web.archive.org/web/20151229212607/http://cbs.umn.edu/sites/cbs.umn.edu/files/public/downloads/When_lions_ruled_France.pdf |archive-date=29 December 2015 |url-status=live}}</ref>
*[[American lion|''P. atrox'']], or the '''American lion''', ranged in the Americas from Canada to possibly [[Patagonia]] during the Late Pleistocene.<ref name="Chimento2017">{{Cite journal |last1=Chimento |first1=N. R. |last2=Agnolin |first2=F. L.  |year=2017 |title=The fossil American lion (''Panthera atrox'') in South America: Palaeobiogeographical implications |journal=Comptes Rendus Palevol |volume=16 |issue=8 |pages=850–864 |doi=10.1016/j.crpv.2017.06.009 |bibcode=2017CRPal..16..850C |url=https://www.researchgate.net/publication/321056731|doi-access=free |hdl=11336/65990 |hdl-access=free }}</ref> It diverged from the cave lion around 165,000 years ago.<ref>{{Cite journal |last1=Salis |first1=Alexander T. |last2=Bray |first2=Sarah C. E. |last3=Lee |first3=Michael S. Y. |last4=Heiniger |first4=Holly |last5=Barnett |first5=Ross |last6=Burns |first6=James A. |last7=Doronichev |first7=Vladimir |last8=Fedje |first8=Daryl |last9=Golovanova |first9=Liubov |last10=Harington |first10=C. Richard |last11=Hockett |first11=Bryan |last12=Kosintsev |first12=Pavel |last13=Lai |first13=Xulong |last14=Mackie |first14=Quentin |last15=Vasiliev |first15=Sergei |date=December 2022 |title=Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge |url=https://onlinelibrary.wiley.com/doi/10.1111/mec.16267 |journal=Molecular Ecology |language=en |volume=31 |issue=24 |pages=6407–6421 |bibcode=2022MolEc..31.6407S |doi=10.1111/mec.16267 |issn=0962-1083 |pmid=34748674 |hdl-access=free |hdl=11343/299180}}</ref> A fossil from [[Edmonton]] dates to 11,355 ± 55 years ago.<ref name="King2013">{{Cite journal |doi=10.1080/08912963.2013.861462 |title=Phylogenetics of ''Panthera'', including ''Panthera atrox'', based on craniodental characters |journal=[[Historical Biology]] |volume=26 |issue=6 |pages=827–833 |url=https://www.researchgate.net/publication/265790587 |date=2014 |last1=King |first1=L. M. |last2=Wallace |first2=S. C. |bibcode=2014HBio...26..827K |s2cid=84229141 }}</ref>

===Evolution===
{{multiple image|align=right|direction=vertical|image1=Panthera Atrox.jpg |thumb|caption1=Skull of an American lion on display at the [[National Museum of Natural History]]|
image2=Cave lion range.png|caption2=red ''[[Panthera spelaea]]''{{break}}blue ''[[Panthera atrox]]''{{break}}green ''Panthera leo''{{break}}{{break}}Maximal range of the modern lion{{break}}and its prehistoric relatives{{break}}in the late Pleistocene}}

The ''Panthera'' [[Lineage (evolution)|lineage]] is estimated to have [[Genetic divergence|genetically diverged]] from the [[common ancestor]] of the [[Felidae]] around {{Ma|9.32|4.47|million years ago}} to {{Ma|11.75|0.97|million years ago}}.<ref name="Johnson2006">{{cite journal |last1=Johnson |first1=W. E. |last2=Eizirik |first2=E. |last3=Pecon-Slattery |first3=J. |last4=Murphy |first4=W. J. |last5=Antunes |first5=A. |last6=Teeling |first6=E. |last7=O'Brien |first7=S. J.  |title=The late miocene radiation of modern Felidae: A genetic assessment |journal=[[Science (journal)|Science]] |volume=311 |issue=5757 |pages=73–77 |date=2006 |pmid=16400146 |doi=10.1126/science.1122277 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=22 August 2020 |archive-date=4 October 2020 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live }}</ref><ref name="Werdelin2010">{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J.  |chapter=Phylogeny and evolution of cats (Felidae) |date=2010 |pages=59–82 |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |chapter-url=https://www.researchgate.net/publication/266755142 |editor1-last=Macdonald |editor1-first=D. W. |editor2-last=Loveridge |editor2-first=A. J. |title=Biology and Conservation of Wild Felids |access-date=10 February 2019 |archive-date=25 September 2018 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}</ref><ref name=Li_al2016>{{cite journal |last1=Li |first1=G. |last2=Davis |first2=B. W. |last3=Eizirik |first3=E. |last4=Murphy |first4=W. J.  |date=2016 |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |volume=26 |issue=1 |pages=1–11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}</ref> Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a [[sister group]] with the [[jaguar]] that diverged {{Ma|3.46|1.22|million years ago}},<ref name=Johnson2006/> but also with the [[leopard]] that diverged {{Ma|3.1|1.95|million years ago}}<ref name="davis2010">{{cite journal |last1=Davis |first1=B. W. |last2=Li |first2=G. |last3=Murphy |first3=W. J.  |title=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, ''Panthera'' (Carnivora: Felidae) |journal=Molecular Phylogenetics and Evolution |year=2010 |volume=56 |issue=1 |pages=64–76 |doi=10.1016/j.ympev.2010.01.036 |pmid=20138224 |bibcode=2010MolPE..56...64D |url=https://www.academia.edu/12157986 |access-date=25 August 2019 |archive-date=21 November 2021 |archive-url=https://web.archive.org/web/20211121100453/https://www.academia.edu/12157986 |url-status=live }}</ref><ref name="mazak2011">{{cite journal |last1=Mazák|first1=J. H. |last2=Christiansen |first2=P. |last3=Kitchener |first3=A. C. |last4=Goswami |first4=A.  |title=Oldest known pantherine skull and evolution of the tiger |journal=PLOS ONE |year=2011 |volume=6 |issue=10 |page=e25483 |doi=10.1371/journal.pone.0025483 |pmid=22016768 |pmc=3189913|bibcode=2011PLoSO...625483M|doi-access=free }}</ref> to {{Ma|4.32|0.02|million years ago}}. [[Hybrid (biology)|Hybridisation]] between lion and [[snow leopard]] ancestors possibly continued until about 2.1 million years ago.<ref name=Li_al2016/> The lion-leopard clade was distributed in the Asian and African [[Palearctic]] since at least the early [[Pliocene]].<ref name=Tseng2014>{{cite journal |author1=Tseng, Z. J. |author2=Wang, X. |author3=Slater, G. J.  |author4=Takeuchi, G. T. |author5=Li, Q. |author6=Liu, J. |author7=Xie, G. |date=2014 |title=Himalayan fossils of the oldest known pantherine establish ancient origin of big cats |journal=Proceedings of the Royal Society B: Biological Sciences |volume=281 |issue=1774 |page=20132686 |doi=10.1098/rspb.2013.2686|pmid=24225466 |pmc=3843846 |doi-access=free }}</ref> The earliest fossils recognisable as lions were found at [[Olduvai Gorge]] in Tanzania and are estimated to be up to 2 million years old.<ref name="Werdelin2010"/>

Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on [[mutation rate]] per generation time of the modern lion. There is no evidence for [[gene flow]] between the two lineages, indicating that they did not share the same geographic area.<ref name="DeManuel_al2020" /> The Eurasian and American cave lions became extinct at the end of the [[last glacial period]] without [[Mitochondrion|mitochondrial]] descendants on other continents.<ref name="BurgerJ-Molecular-phylogeny"/><ref name=Barnett>{{Cite journal |last1=Barnett |first1=R. |last2=Shapiro |first2=B. |author-link2=Beth Shapiro |last3=Barnes |first3=I. |last4=Ho |first4=S. Y. W. |last5=Burger |first5=J. |author-link5=Joachim Burger |last6=Yamaguchi |first6=N. |last7=Higham |first7=T. F. G. |last8=Wheeler |first8=H. T. |title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=[[Molecular Ecology]] |date=2009 |volume=18 |issue=8 |pages=1668–1677 |pmid=19302360 |doi=10.1111/j.1365-294X.2009.04134.x |first9=W. |last9=Rosendahl |last10=Sher |first10=A. V. |last11=Sotnikova |first11=M. |last12=Kuznetsova |first12=T. |last13=Baryshnikov |first13=G. F. |last14=Martin |first14=L. D. |last15=Harington |first15=C. R. |last16=Burns |first16=J. A. |last17=Cooper |first17=A. |bibcode=2009MolEc..18.1668B |s2cid=46716748  |url=http://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |archive-url=https://web.archive.org/web/20170808225555/http://www.zin.ru/labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |archive-date=8 August 2017 |url-status=live}}</ref><ref>{{cite journal |author1=Argant, A. |author2=Brugal, J.-P.  |year=2017|title=The cave lion ''Panthera (Leo) spelaea'' and its evolution: ''Panthera spelaea intermedia'' nov. subspecies |journal=Acta Zoologica Cracoviensia |volume=60 |issue=2 |pages=58–103 |doi=10.3409/azc.60_2.59 |doi-access=free}}</ref> The modern lion was probably widely distributed in Africa during the [[Middle Pleistocene]] and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago.<ref name=Barnett_al2014>{{cite journal |author1=Barnett, R. |author2=Yamaguchi, N. |author3=Shapiro, B. |author4=Ho, S. Y. |author5=Barnes, I. |author6=Sabin, R. |author7=Werdelin, L. |author8=Cuisin, J.  |author9=Larson, G. |year=2014 |title=Revealing the maternal demographic history of ''Panthera leo'' using ancient DNA and a spatially explicit genealogical analysis |journal=BMC Evolutionary Biology |volume=14 |issue=1 |page=70 |doi=10.1186/1471-2148-14-70|pmid=24690312 |pmc=3997813 |doi-access=free |bibcode=2014BMCEE..14...70B }}</ref> They shared a common ancestor probably between 98,000 and 52,000 years ago.<ref name="DeManuel_al2020" /> Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago.<ref name=Barnett_al2014/>

Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous [[mutation]]s in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.<ref name=Bertola2011>{{Cite journal |last1=Bertola |first1=L. D. |last2=Van Hooft |first2=W. F. |last3=Vrieling |first3=K. |last4=Uit De Weerd |first4=D. R. |last5=York |first5=D. S. |last6=Bauer |first6=H. |last7=Prins |first7=H. H. T. |last8=Funston |first8=P. J. |last9=Udo De Haes |first9=H. A. |last10=Leirs |first10=H. |last11=Van Haeringen |first11=W. A. |last12=Sogbohossou |first12=E. |last13=Tumenta |first13=P. N. |last14=De Iongh |first14=H. H.  |year=2011 |title=Genetic diversity, evolutionary history and implications for conservation of the lion (''Panthera leo'') in West and Central Africa |journal=Journal of Biogeography |volume=38 |issue=7 |pages=1356–1367 |doi=10.1111/j.1365-2699.2011.02500.x|bibcode=2011JBiog..38.1356B |s2cid=82728679 }}</ref> A whole genome-wide sequence of lion samples showed that samples from West Africa shared [[alleles]] with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in the [[Nile Basin]] during the early [[Holocene]].<ref name="DeManuel_al2020" />

===Hybrids===

{{Further|Panthera hybrid}}

In zoos, lions have been bred with [[tiger]]s to create hybrids for the curiosity of visitors or for scientific purpose.<ref>{{cite journal |author=Pocock, R. I. |year=1898 |title=Lion-Tiger Hybrid |journal=Nature |volume=58 |issue=1496 |page=200 |doi=10.1038/058200b0 |url=https://zenodo.org/record/1889713 |bibcode=1898Natur..58Q.200P |s2cid=4056029 |doi-access=free |access-date=7 December 2019 |archive-date=26 March 2022 |archive-url=https://web.archive.org/web/20220326042949/https://zenodo.org/record/1889713 |url-status=live }}</ref><ref>{{cite book |author=Benirschke, K. |title=Comparative Aspects of Reproductive Failure |chapter=Sterility and Fertility of Interspecific Mammalian Hybrids |pages=218–234 |publisher=Springer |location=Berlin, Heidelberg |year=1967 |doi=10.1007/978-3-642-48949-5_12 |isbn=978-3-642-48949-5}}</ref> The [[liger]] is bigger than a lion and a tiger, whereas most [[tigon]]s are relatively small compared to their parents because of reciprocal gene effects.<ref name="shi">{{cite thesis |last=Shi |first=W. |year=2005 |title=Growth and Behaviour: Epigenetic and Genetic Factors Involved in Hybrid Dysgenesis |type=PhD |series=Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology |publisher=Acta Universitatis Upsaliensis |location=Uppsala |chapter=Hybrid dysgenesis effects |pages=8–10 |chapter-url=http://uu.diva-portal.org/smash/get/diva2:165749/FULLTEXT01.pdf |archive-url=https://web.archive.org/web/20190518050521/http://uu.diva-portal.org/smash/get/diva2:165749/FULLTEXT01.pdf |archive-date=18 May 2019 |url-status=live|url=http://urn.kb.se/resolve?urn=urn:nbn:se:uu:diva-4784 }}</ref><ref name="Carnivores">{{Cite book |last1=Rafferty |first1=J. P. |title=Carnivores: Meat-eating Mammals |date=2011 |location=New York |publisher=The Rosen Publishing Group |isbn=978-1-61530-340-3 |chapter=The Liger |page=120 |chapter-url=https://books.google.com/books?id=EMui7zVOqeUC&pg=PA120 |access-date=4 July 2014 |archive-date=5 May 2024 |archive-url=https://web.archive.org/web/20240505150613/https://books.google.com/books?id=EMui7zVOqeUC&pg=PA120#v=onepage&q&f=false |url-status=live }}</ref> The [[leopon]] is a hybrid between a lion and leopard.<ref>{{Cite journal |author1=Zhang, Z. |author2=Chen, J. |author3=Li, L. |author4=Tao, M. |author5=Zhang, C. |author6=Qin, Q. |author7=Xiao, J. |author8=Liu, Y. |author9=Liu, S. |title=Research advances in animal distant hybridization |year=2014 |journal=Science China Life Sciences |volume=57 |issue=9 |pages=889–902 |doi=10.1007/s11427-014-4707-1 |pmid=25091377 |s2cid=18179301 |url=https://link.springer.com/content/pdf/10.1007/s11427-014-4707-1.pdf |archive-url=https://web.archive.org/web/20181030104852/https://link.springer.com/content/pdf/10.1007%2Fs11427-014-4707-1.pdf |archive-date=30 October 2018 |url-status=live|doi-access=free }}</ref>