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==Evolution== The geographic origin of the genus ''Panthera'' is uncertain, though the earliest known definitive species ''[[Panthera principialis]]'' is from [[Tanzania]].<ref name=principialis/> ''P. blytheae'' from northern [[Central Asia]], originally described as the oldest known ''Panthera'' species, is suggested to be similar in skull features to the snow leopard,<ref name=Tseng_al2014/> but subsequent studies have since agreed that it is not a member of or a related species of the snow leopard lineage and that it belongs to a different genus ''[[Palaeopanthera]]''.<ref>{{cite journal |author1=Geraads, D. |author2=PeignΓ©, S |title=Re-appraisal of ''Felis pamiri'' Ozansoy 1959 (Carnivora, Felidae) from the upper Miocene of Turkey: the earliest pantherine cat? |journal=Journal of Mammalian Evolution |volume=24 |issue=4 |pages=415β425 |year=2017 |url=https://hal.sorbonne-universite.fr/hal-01675275/document |doi=10.1007/s10914-016-9349-6|s2cid=207195894}}</ref><ref>{{cite journal|last1=Hemmer|first1=H. |date=2023 |title=The evolution of the palaeopantherine cats, ''Palaeopanthera'' gen. nov. ''blytheae'' (Tseng et al., 2014) and ''Palaeopanthera pamiri'' (Ozansoy, 1959) comb. nov. (Mammalia, Carnivora, Felidae) |journal=Palaeobiodiversity and Palaeoenvironments |volume=103 |issue=4 |pages=827β839 |doi=10.1007/s12549-023-00571-5 |bibcode=2023PdPe..103..827H |s2cid=257842190}}</ref><ref>{{Cite journal|last1=Jiangzuo |first1=Q. |last2=Madurell-Malapeira |first2=J. |last3=Li |first3=X. |last4=Estraviz-LΓ³pez |first4=D. |last5=Mateus |first5=O. |last6=Testu |first6=A. |last7=Li |first7=S. |last8=Wang |first8=S. |last9=Deng |first9=T. |title=Insights on the evolution and adaptation toward high-altitude and cold environments in the snow leopard lineage |year=2025 |journal=Science Advances |volume=11 |issue=3 |pages=eadp5243 |doi=10.1126/sciadv.adp5243 |pmid=39813339 |pmc=11734717 |doi-access=free }}</ref> The tiger, snow leopard, and [[clouded leopard]] [[genetic lineage]]s likely dispersed in Southeast Asia during the [[Late Miocene]].<ref name=Tseng_al2014>{{cite journal |last1=Tseng |first1=Z.J. |last2=Wang |first2=X. |last3=Slater |first3=G.J. |last4=Takeuchi |first4=G.T. |last5=Li |first5=Q. |last6=Liu |first6=J. |last7=Xie |first7=G. |name-list-style=amp |year=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}}</ref> Genetic studies indicate that the [[Pantherinae|pantherine cats]] diverged from the subfamily [[Felinae]] between six and ten 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. |name-list-style=amp |year=2006 |doi=10.1126/science.1122277 |title=The Late Miocene radiation of modern Felidae: A genetic assessment |journal=Science |volume=311 |pages=73β77 |pmid=16400146 |issue=5757 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=28 June 2019 |archive-date=4 October 2020 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live}}</ref> The genus ''[[Neofelis]]'' is [[sister group|sister]] to ''Panthera''.<ref name=Johnson2006/><ref name=Janczewski>{{cite journal |last1=Janczewski |first1=D.N. |last2=Modi |first2=W.S. |last3=Stephens |first3=J.C. |last4=O'Brien |first4=S.J. |name-list-style=amp |year=1996 |title=Molecular evolution of mitochondrial 12S RNA and cytochrome b sequences in the pantherine lineage of ''Felidae'' |journal=Molecular Biology and Evolution |volume=12 |issue=4 |pages=690β707 |doi=10.1093/oxfordjournals.molbev.a040232 |pmid=7544865 |doi-access=free}}</ref><ref name=Johnson1997>{{cite journal |last1=Johnson |first1=W. E. |last2=O'Brien |first2=S.J. |name-list-style=amp |year=1997 |title=Phylogenetic reconstruction of the ''Felidae'' using 16S rRNA and NADH-5 mitochondrial genes |journal=Journal of Molecular Evolution |volume=44 |issue=S1 |pages=S98βS116 |doi=10.1007/PL00000060 |pmid=9071018 |bibcode=1997JMolE..44S..98J |s2cid=40185850 |url=https://zenodo.org/record/1232587 |access-date=28 June 2019 |archive-date=4 October 2020 |archive-url=https://web.archive.org/web/20201004075723/https://zenodo.org/record/1232587 |url-status=live}}</ref><ref name=Yu1>{{cite journal |last1=Yu |first1=L. |last2=Zhang |first2=Y.P. |name-list-style=amp |year=2005 |title=Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores |journal=Molecular Phylogenetics and Evolution |volume=35 |issue=2 |pages=483β495 |doi=10.1016/j.ympev.2005.01.017 |pmid=15804417|bibcode=2005MolPE..35..483Y }}</ref> The clouded leopard appears to have [[Genetic divergence|diverged]] about {{Ma|8.66}}. ''Panthera'' diverged from other cat species about {{Ma|11.3}} and then evolved into the species tiger about {{Ma|6.55}}, snow leopard about {{Ma|4.63}} and leopard about {{Ma|4.35}}. Mitochondrial sequence data from fossils suggest that the [[American lion]] (''P. atrox'') is a sister lineage to ''[[Panthera spelaea]]'' (the Eurasian cave or steppe lion) that diverged about {{Ma|0.34}}, and that both ''P. atrox'' and ''P. spelaea'' are most closely related to lions among living ''Panthera'' species.<ref name="Barnett, 2009">{{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. |last9=Rosendahl |first9=W. |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. |name-list-style=amp |title=Phylogeography of lions (''Panthera leo'' ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=[[Molecular Ecology]] |year=2009 |volume=18 |issue=8 |pages=1668β1677 |pmid=19302360 |doi=10.1111/j.1365-294X.2009.04134.x |bibcode=2009MolEc..18.1668B |s2cid=46716748 |url=https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |access-date=24 February 2019 |archive-date=8 August 2017 |archive-url=https://web.archive.org/web/20170808225555/https://www.zin.ru/Labs/theriology/eng/staff/baryshnikov/references/barnett_et_al_2009.pdf |url-status=live}}</ref> The snow leopard is nested within ''Panthera'' and is the [[sister species]] of the tiger.<ref name=Davis2010>{{cite journal |last1=Davis |first1=B.W. |title=Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, ''Panthera'' (''Carnivora'': ''Felidae'') |author2=Li, G. |author3=Murphy, W.J. |journal=Molecular Phylogenetics and Evolution |volume=56 |date=2010 |issue=1 |pages=64β76 |pmid=20138224 |doi=10.1016/j.ympev.2010.01.036|bibcode=2010MolPE..56...64D }}</ref> Results of a 2016 study based on analysis of biparental [[nuclear genome]]s suggest the following relationships of living ''Panthera'' species:<ref name=Li_al2016>{{cite journal |author=Li, G. |author2=Davis, B. W. |author3=Eizirik, E. |name-list-style=amp |author4=Murphy, W. J. |year=2016 |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |volume=26 |issue=1 |pages=1β11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}</ref> The extinct species ''[[Panthera gombaszoegensis]]'', was probably closely related to the modern jaguar. The first fossil remains were [[Excavation (archaeology)|excavated]] in [[Olivola]], in Italy, and date to {{Ma|1.6}}.<ref>{{cite journal |last1=Hemmer |first1=H. |last2=Kahlke |first2=R.D. |last3=Vekua |first3=A.K. |name-list-style=amp |year=2001 |title=The Jaguar β ''Panthera onca gombaszoegensis'' (Kretzoi, 1938) (Carnivora: Felidae) in the late lower Pleistocene of Akhalkalaki (south Georgia; Transcaucasia) and its evolutionary and ecological significance | journal=Geobios |volume=34 |issue=4 |pages=475β486 |doi=10.1016/s0016-6995(01)80011-5 |bibcode=2001Geobi..34..475H}}</ref> Fossil remains found in South Africa that appear to belong within the ''Panthera'' lineage date to about {{Ma|2.0|3.8}}.<ref>{{cite journal |last1=Turner |first1=A. |year=1987 |title=New fossil carnivore remains from the Sterkfontein hominid site (''Mammalia'': ''Carnivora'') |journal=Annals of the Transvaal Museum |volume=34 |issue=15 |pages=319β347 |url=https://journals.co.za/content/nfi_annalstm/34/15/AJA00411752_121?crawler=true&mimetype=application/pdf}}</ref>
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