Editing Leopard (section)

===Evolution===
[[File:Two cladograms for Panthera.svg|thumb|upright|Two cladograms proposed for ''Panthera''. The upper cladogram is based on the 2006<ref name=johnson2006/> and 2009<ref name=werdelin2009/> studies, while the lower is based on the 2010<ref name=davis2010/> and 2011<ref name="mazak2011"/> studies.]]
Results of [[phylogenetic]] studies based on [[nuclear DNA]] and [[mitochondrial DNA]] analysis showed that the last [[common ancestor]] of the ''Panthera'' and ''[[Neofelis]]'' genera is thought to have lived about {{Ma|6.37}}. ''Neofelis'' [[Genetic divergence|diverged]] about {{Ma|8.66}} from the ''Panthera'' [[Genetic lineage|lineage]]. The tiger diverged about {{Ma|6.55}}, followed by the snow leopard about {{Ma|4.63}} and the leopard about {{Ma|4.35}}. The leopard is a [[sister taxon]] to a [[clade]] within ''Panthera'', consisting of the lion and the jaguar.<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 |year=2006 |volume=311 |issue=5757 |pages=73–77 |name-list-style=amp |pmid=16400146 |doi=10.1126/science.1122277 |bibcode=2006Sci...311...73J |s2cid=41672825 |url=https://zenodo.org/record/1230866 |access-date=2019-07-05 |archive-date=2020-10-04 |archive-url=https://web.archive.org/web/20201004075725/https://zenodo.org/record/1230866 |url-status=live }}</ref><ref name=werdelin2009>{{cite book |last1=Werdelin |first1=L. |last2=Yamaguchi |first2=N. |last3=Johnson |first3=W. E. |last4=O'Brien |first4=S. J. |name-list-style=amp |chapter=Phylogeny and evolution of cats (Felidae) |year=2010 |pages=59–82 |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 |publisher=Oxford University Press |location=Oxford, UK |isbn=978-0-19-923445-5 |access-date=2018-09-25 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925141956/https://www.researchgate.net/publication/266755142 |url-status=live }}</ref>

Results of a phylogenetic analysis of chemical [[secretion]]s amongst cats indicated that the leopard is closely related to the lion.<ref>{{cite journal |last1=Bininda-Emonds |first1=O. R. P. |last2=Decker-Flum |first2=D. M. |last3=Gittleman |first3=J. L. |name-list-style=amp |title=The utility of chemical signals as phylogenetic characters: an example from the Felidae |journal=Biological Journal of the Linnean Society |year=2001 |volume=72 |issue=1 |pages=1–15 |doi=10.1111/j.1095-8312.2001.tb01297.x |doi-access=free}}</ref>
The geographic origin of the ''Panthera'' is most likely northern Central Asia. The leopard-lion clade was distributed in the Asian and African [[Palearctic]] since at least the early [[Pliocene]].<ref>{{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. |name-list-style=amp |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 }}</ref> The leopard-lion clade diverged 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. |name-list-style=amp |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=http://web.csulb.edu/~acarter3/course-evolution/files/cat-phylogeny.pdf |url-status=dead |archive-url=https://web.archive.org/web/20160305131606/http://web.csulb.edu/~acarter3/course-evolution/files/cat-phylogeny.pdf |archive-date=2016-03-05}}</ref><ref name=mazak2011>{{cite journal |last1=Mazák |first1=J. H. |last2=Christiansen |first2=P. |last3=Kitchener|first3=A. C. |last4=Goswami |first4=A. |name-list-style=amp |title=Oldest known pantherine skull and evolution of the tiger |journal=PLOS ONE |year=2011 |volume=6 |issue=10 |pages=e25483 |doi=10.1371/journal.pone.0025483 |pmid=22016768 |pmc=3189913 |bibcode=2011PLoSO...625483M |doi-access=free}}</ref> Additionally, a 2016 study revealed that the [[mitochondrial genome]]s of the leopard, lion and [[snow leopard]] are more similar to each other than their [[nuclear genome]]s, indicating that their ancestors [[Hybridization (biology)|hybridized]] with the snow leopard at some point in their [[evolution]].<ref>{{cite journal |last1=Li |first1=G. |last2=Davis |first2=B. W. |last3=Eizirik |first3=E. |last4=Murphy |first4=W. J. |name-list-style=amp |title=Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae) |journal=Genome Research |date=2016 |volume=26 |issue=1 |pages=1–11 |doi=10.1101/gr.186668.114 |pmid=26518481 |pmc=4691742}}</ref>

The oldest unambiguous [[fossil]]s of the leopard are from Eastern Africa, dating to around 2 million years ago.<ref name=Paijmans>{{Cite journal |last1=Paijmans |first1=J. L. A. |last2=Barlow |first2=A. |last3=Förster |first3=D. W. |last4=Henneberger |first4=K. |last5=Meyer |first5=M. |last6=Nickel |first6=B. |last7=Nagel |first7=D. |last8=Havmøller |first8=R. W. |last9=Baryshnikov |first9=G. F. |last10=Joger |first10=U. |last11=Rosendahl |first11=W. |last12=Hofreiter |first12=M. |name-list-style=amp |date=2018 |title=Historical biogeography of the leopard (''Panthera pardus'') and its extinct Eurasian populations |journal=BMC Evolutionary Biology |volume=18 |issue=1 |page=156 |doi=10.1186/s12862-018-1268-0 |pmc=6198532 |pmid=30348080 |doi-access=free |bibcode=2018BMCEE..18..156P}}</ref>

Leopard-like fossil bones and teeth possibly dating to the [[Pliocene]] were excavated in [[Perrier, Puy-de-Dôme|Perrier]] in [[France]], northeast of [[London]], and in [[Valdarno]], [[Italy]]. Until 1940, similar fossils dating back to the [[Pleistocene]] were excavated mostly in [[loess]] and caves at 40 sites in Europe, including [[Furninha]] Cave near [[Lisbon]], [[Genista Caves]] in [[Gibraltar]], and [[Santander Province]] in northern [[Spain]] to several sites across France, [[Switzerland]], Italy, [[Austria]], [[Germany]], in the north up to [[Derby]] in [[England]], in the east to [[Přerov]] in the [[Czech Republic]] and the [[Baranya (region)|Baranya]] in southern [[Hungary]].<ref name=Schmid1940>{{cite journal |last=Schmid |first= E. |year=1940 |title=Variationstatistische Untersuchungen am Gebiss pleistozäner und rezenter Leoparden und anderer Feliden |journal=Zeitschrift für Säugetierkunde |volume=15 |pages=1–179 |url=https://archive.org/details/zeitschrift1519401943deut/page/n11}}</ref>
Leopards arrived in Eurasia during the late Early to Middle Pleistocene around 1.2<ref name=Marciszak>{{Cite journal |last1=Marciszak |first1=A. |last2=Lipecki |first2=G. |last3=Gornig |first3=W. |last4=Matyaszczyk |first4=L. |last5=Oszczepalińska |first5=O. |last6=Nowakowski |first6=D. |last7=Talamo |first7=S. |title=The first radiocarbon-dated remains of the Leopard ''Panthera pardus'' (Linnaeus, 1758) from the Pleistocene of Poland |date=2022 |journal=[[Radiocarbon (journal)|Radiocarbon]] |volume=64 |issue=6 |pages=1359–1372 |doi=10.1017/RDC.2022.33 |bibcode=2022Radcb..64.1359M |s2cid=255850069 |hdl=11585/887180 |hdl-access=free |url=https://www.cambridge.org/core/product/identifier/S0033822222000339/type/journal_article |access-date=2023-07-18 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145527/https://www.cambridge.org/core/journals/radiocarbon/article/first-radiocarbondated-remains-of-the-leopard-panthera-pardus-linnaeus-1758-from-the-pleistocene-of-poland/14B7D49EB6F90245554DCBA441123E3B |url-status=live }}</ref> to 0.6 million years ago.<ref name=Paijmans />
Four European Pleistocene leopard subspecies were proposed. ''P. p. begoueni'' from the beginning of the [[Early Pleistocene]] was replaced about {{Ma|0.6}} by ''P. p. sickenbergi'', which in turn was replaced by ''P. p. antiqua'' around 0.3 million years ago.<ref name=Diedrich2013/> [[Panthera pardus spelaea|''P. p. spelaea'']] is the most recent subspecies that appeared at the beginning of the [[Late Pleistocene]] and survived until about 11,000 years ago and possibly into the early [[Holocene]] in the [[Iberian Peninsula]].<ref name=Diedrich2013>{{cite journal |last1=Diedrich|first1=C. G. |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |year=2013 |volume=76 |pages=167–193 |doi=10.1016/j.quascirev.2013.05.009 |bibcode=2013QSRv...76..167D}}</ref><ref>{{Cite journal |last1=Sauqué |first1=V. |name-list-style=amp |last2=Cuenca‑Bescós |first2=G. |date=2013 |title=The Iberian Peninsula, the last European refugium of ''Panthera pardus'' Linnaeus 1758 during the Upper Pleistocene |journal=Quaternaire |url=https://journals.openedition.org/quaternaire/6468 |volume=24 |issue=1 |pages=13–24 |doi=10.4000/quaternaire.6468 |access-date=2023-07-18 |archive-date=2023-07-18 |archive-url=https://web.archive.org/web/20230718210435/https://journals.openedition.org/quaternaire/6468 |url-status=live |url-access=subscription }}</ref>

Leopards depicted in cave paintings in [[Chauvet Cave]] provide indirect evidence of leopard presence in Europe.<ref name=Diedrich2013/>
Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central [[Poland]].<ref>{{Cite journal |last1=Marciszak |first1=A. |last2=Stefaniak |first2=K. |name-list-style=amp |date=2010 |title=Two forms of cave lion: Middle Pleistocene ''Panthera spelaea fossilis'' Reichenau, 1906 and Upper Pleistocene ''Panthera spelaea spelaea'' Goldfuss, 1810 from the Bísnik Cave, Poland |url=https://www.researchgate.net/publication/233669138 |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117 |access-date=2017-08-24 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925142142/https://www.researchgate.net/publication/233669138 |url-status=live }}</ref><ref name=Marciszak/>
Fossil remains were also excavated in the Iberian<ref>{{Cite journal |last1=Sauqué |first1=V. |last2=Rabal-Garcés |first2=R. |last3=Cuenca-Bescós |first3=G. |name-list-style=amp |date=2016 |title=Carnivores from Los Rincones, a leopard den in the highest mountain of the Iberian range (Moncayo, Zaragoza, Spain) |journal=[[Historical Biology]] |volume=28 |issue=4 |pages=479–506 |doi=10.1080/08912963.2014.977882 |doi-access=free |bibcode=2016HBio...28..479S}}</ref><ref>{{Cite journal |last1=Sanchis |first1=A. |last2=Real |first2=C. |last3=Sauqué |first3=V. |last4=Núñez-Lahuerta |first4=C. |last5=Égüez |first5=N. |last6=Tormo |first6=C. |last7=Ripoll |first7=M. P. |last8=Carrión |first8=Y. M. |last9=Duarte |first9=E. |last10=de la Rasilla |first10=M. |name-list-style=amp |date=2019 |title=Neanderthal and carnivore activities at Llonin Cave, Asturias, northern Iberian Peninsula: Faunal study of Mousterian levels (MIS 3) |journal=[[Comptes Rendus Palevol]] |volume=18 |issue=1 |pages=113–141 |bibcode=2019CRPal..18..113S |doi=10.1016/j.crpv.2018.06.001 |s2cid=134157124 |hdl-access=free |hdl=10651/49880}}</ref> and [[Italian Peninsula]],<ref>{{Cite journal |last1=Ghezzo |first1=E. |last2=Rook |first2=L. |date=2015 |name-list-style=amp |title=The remarkable ''Panthera pardus'' (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology |journal=[[Quaternary Science Reviews]] |volume=110 |pages=131–151 |doi=10.1016/j.quascirev.2014.12.020}}</ref><ref>{{Cite journal |last=Mecozzi |first=Beniamino |last2=Sardella |first2=Raffaele |last3=Boscaini |first3=Alberto |last4=Cherin |first4=Marco |last5=Costeur |first5=Loïc |last6=Madurell-Malapeira |first6=Joan |last7=Pavia |first7=Marco |last8=Profico |first8=Antonio |last9=Iurino |first9=Dawid A. |date=15 June 2021 |title=The tale of a short-tailed cat: New outstanding Late Pleistocene fossils of Lynx pardinus from southern Italy |url=https://www.sciencedirect.com/science/article/pii/S0277379121000470 |journal=[[Quaternary Science Reviews]] |language=en |volume=262 |pages=106840 |doi=10.1016/j.quascirev.2021.106840 |access-date=7 November 2024 |via=Elsevier Science Direct|hdl=2434/959858 |hdl-access=free }}</ref> and in the [[Balkans]].<ref>{{Cite journal |last1=Sommer |first1=R. S. |last2=Benecke |first2=N. |name-list-style=amp |date=2006 |title=Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review |journal=[[Journal of Zoology]] |volume=269 |issue=1 |pages=7–19 |doi=10.1111/j.1469-7998.2005.00040.x}}</ref><ref>{{Cite journal |last1=Miracle |first1=P. T. |last2=Lenardić |first2=J. M. |last3=Brajković |first3=D. |date=2010 |title=Last glacial climates, "Refugia", and faunal change in Southeastern Europe: Mammalian assemblages from Veternica, Velika pećina, and Vindija caves (Croatia) |journal=[[Quaternary International]] |volume=212 |issue=2 |pages=137–148 |doi=10.1016/j.quaint.2009.06.003|bibcode=2010QuInt.212..137M }}</ref>
Leopard fossils dating to the Pleistocene were also excavated in the [[Japanese archipelago]].<ref name=JapanMammals>{{cite book |author=Izawa, M. |author2=Nakanishi, N. |name-list-style=amp |chapter=Felidae |pages=226−231 |editor=Ohdachi, S. D. |editor2=Ishibashi, Y. |editor3=Iwasa, M. A. |editor4=Saitoh, T. |year=2015 |title=The Wild Mammals of Japan |publisher=Shoukadoh Book Sellers and the Mammalogical Society of Japan |edition=Second |isbn=978-4-87974-691-7 |location=Kyoto}}</ref> Leopard fossils were also found in [[Taiwan]].<ref>{{cite journal |author1=Chi T.-C. |name-list-style=amp |author2=Gan Y. |author3=Yang T.-R. |author4=Chang, C.-H. |year=2021 |title=First report of leopard fossils from a limestone cave in Kenting area, southern Taiwan |journal=PeerJ |volume=9 |page=e12020 |doi=10.7717/peerj.12020 |pmid=34513335 |pmc=8388558 |doi-access=free}}</ref>