Editing Lion (section)

===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" />