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===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>
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