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{{Short description|Species of cat native to Africa and Asia}}
{{Other uses|Leopard (disambiguation)|Leopards (disambiguation)}}
{{Pp-semi-indef}}
{{Pp-move}}
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{{Speciesbox
| name = Leopard
| fossil_range = {{fossilrange|Early Pleistocene | Present}}
| image = African leopard male (cropped).jpg <!--- Please do not change consensus image. This was agreed upon at the talk page ---->
| image_caption = Male [[African leopard]] in [[Maasai Mara National Reserve]], [[Kenya]]
| status = VU
| status_system = IUCN3.1
| status_ref = <ref name=IUCN>{{cite iucn |title=''Panthera pardus'' |name-list-style=amp |author1=Stein, A.B. |author2=Gerngross, P. |author3=Al Hikmani, H. |author4=Balme, G. |author5=Bertola, L. |author6=Drouilly, M. |author7=Farhadinia, M.S. |author8=Feng, L. |author9=Ghoddousi, A. |author10=Henschel, P. |author11=Jhala, Y. |author12=Khorozyan, I. |author13=Kittle, A. |author14=Laguardia, A. |author15=Luo, S.-J. |author16=Mann, G. |author17=Miquelle, D. |author18=Moheb, Z. |author19=Raza, H. |author20=Rostro-García, S. |author21=Shivakumar, S. |author22=Song, D. |author23=Wibisono, H. |year=2024 |page=e.T15954A254576956 |access-date=30 June 2024}}</ref>
|
| status2_system = CITES
| status2_ref = <ref name=IUCN/>
| taxon = Panthera pardus<ref name=msw3>{{MSW3 Wozencraft |id=14000250 |pages=547 |heading=Species ''Panthera pardus''}}</ref>
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
| subdivision_ranks = Subspecies
| subdivision = See [[#Living subspecies|text]]
| range_map = Leopard distribution.jpg
| range_map_caption = Present and historical distribution of the leopard<ref name=IUCN/>
| synonyms =
}}
The '''leopard''' ('''''Panthera pardus''''') is one of the five extant cat [[species]] in the [[Genus (biology)|genus]] ''[[Panthera]]''. It has a pale yellowish to dark golden fur with dark spots grouped in [[Rosette (zoology)|rosette]]s. Its body is slender and muscular reaching a length of {{cvt|92|-|183|cm}} with a {{cvt|66|-|102|cm|}} long tail and a shoulder height of {{cvt|60|-|70|cm}}. Males typically weigh {{cvt|30.9|-|72|kg}}, and females {{cvt|20.5|-|43|kg}}.
The leopard was first [[Species description|described]] in 1758, and several [[subspecies]] were proposed in the 19th and 20th centuries. Today, eight subspecies are recognised in its wide range in [[Africa]] and [[Asia]]. It initially evolved in Africa during the [[Early Pleistocene]], before migrating into Eurasia around the Early–[[Middle Pleistocene]] transition. [[European leopard|Leopards were formerly present across Europe]], but became extinct in the region at around the end of the [[Late Pleistocene]]-early [[Holocene]].
The leopard is adapted to a variety of habitats ranging from [[rainforest]] to [[steppe]], including [[Aridity|arid]] and montane areas. It is an [[Generalist and specialist species|opportunistic]] predator, hunting mostly [[ungulate]]s and [[primate]]s. It relies on its spotted pattern for [[camouflage]] as it stalks and ambushes its prey, which it sometimes drags up a tree. It is a [[solitary animal]] outside the mating season and when raising cubs. Females usually give birth to a litter of 2–4 cubs once in 15–24 months. Both male and female leopards typically reach [[sexual maturity]] at the age 2–2.5 years.
Listed as [[Vulnerable species|Vulnerable]] on the [[IUCN Red List]], leopard populations are currently threatened by [[habitat loss]] and fragmentation, and are declining in large parts of the global range. Leopards have had cultural roles in [[Ancient Greece]], [[West Africa]] and modern Western culture. [[Animal print|Leopard skins]] are popular in fashion.
==Etymology==
The English name "leopard" comes from [[Old French]] {{lang|fro|leupart}} or [[Middle French]] {{lang|frm|liepart}}, that derives from [[Latin]] {{lang|la|leopardus}} and [[ancient Greek]] {{lang|grc|λεόπαρδος}} ({{Transliteration|grc|leopardos}}). {{Transliteration|grc|Leopardos}} could be a [[Compound (linguistics)|compound]] of {{lang|grc|λέων}} ({{Transliteration|grc|leōn}}), meaning {{gloss|lion}}, and {{lang|grc|πάρδος}} ({{Transliteration|grc|pardos}}), meaning {{gloss|spotted}}.<ref name=lewis>{{cite book |last1=Lewis |first1=C. T. |author1-link=Charlton Thomas Lewis |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |___location=Oxford |publisher=Clarendon Press |chapter=lěǒpardus |page=1069 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1069}}</ref><ref name=liddell>{{cite book |last1=Liddell |first1=H. G. |author1-link=Henry Liddell |last2=Scott |first2=R. |author2-link=Robert Scott (philologist) |name-list-style=amp |year=1889 |title=A Greek–English Lexicon |___location=Oxford |publisher=Clarendon Press |chapter=λέο-πάρδος |page=884 |chapter-url=https://archive.org/details/greekenglishlex00liddrich/page/884}}</ref><ref>{{Cite book |last=Partridge |first=E. |author-link=Eric Partridge |title=Origins: A Short Etymological Dictionary of Modern English |publisher=Greenwich House |year=1983 |___location=New York |isbn=978-0-517-41425-5 |page=349 |url=https://archive.org/details/originsshortetym0000part/page/349}}</ref> The word {{lang|drc|λεόπαρδος}} originally referred to a [[cheetah]] (''Acinonyx jubatus'').<ref>{{Cite journal |last=Nicholas |first=N. |s2cid=56160515 |year=1999|title=A conundrum of cats: pards and their relatives in Byzantium |journal=[[Greek, Roman, and Byzantine Studies]] |volume=40 |pages=253–298}}</ref>
"Panther" is another common name, derived from Latin {{lang|la|panther}} and ancient Greek {{lang|grc|πάνθηρ}} ({{Transliteration|grc|pánthēr}});<ref name=lewis/> The [[Generic name (biology)|generic name]] ''Panthera'' originates in Latin {{lang|la|panthera}}, a hunting net for catching wild beasts to be used by the [[Roman people|Roman]]s in combats.<ref>{{cite book |last1=Lewis |first1=C. T. |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |___location=Oxford |publisher=Clarendon Press |chapter=panthera |page=1298 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1317}}</ref> {{lang|la|Pardus}} is the [[Grammatical gender|masculine singular]] form.<ref>{{cite book |last1=Lewis |first1=C. T. |last2=Short |first2=C. |name-list-style=amp |year=1879 |title=A Latin Dictionary |___location=Oxford |publisher=Clarendon Press |chapter=''pardus'' |pages=1302 |chapter-url=https://archive.org/details/in.ernet.dli.2015.61236/page/n1321}}</ref>
==Taxonomy==
[[File:Panthera pardus subspecies map.png|thumb|Map showing approximate distribution of leopard subspecies]]
''Felis pardus'' was the [[scientific name]] proposed by [[Carl Linnaeus]] in 1758.<ref name="linnaeus">{{cite book |author=Linnaeus, 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 |pages=41−42 |chapter=Felis pardus |chapter-url=https://archive.org/stream/mobot31753000798865#page/41/mode/2up}} {{in lang|la}}</ref>
The [[Generic name (biology)|generic name]] ''[[Panthera]]'' was first used by [[Lorenz Oken]] in 1816, who included all the known spotted cats into this group.<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 |access-date=2019-04-06 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145524/https://books.google.com/books?id=S5o5AAAAcAAJ&pg=PA1052#v=onepage&q&f=false |url-status=live }}</ref>
Oken's classification was not widely accepted, and ''[[Felis]]'' or ''[[Leopardus]]'' was used as the generic name until the early 20th century.<ref name=Ellerman1966>{{cite book|last1=Ellerman |first1=J. R. |last2=Morrison-Scott |first2=T. C. S. |year=1966 |url=https://archive.org/stream/checklistofindia00elle#page/314/mode/2up |title=Checklist of Palaearctic and Indian mammals 1758 to 1946 |edition=Second|publisher=British Museum of Natural History |___location= London |pages=315–317}}</ref>
The leopard was designated as the [[type species]] of ''Panthera'' by [[Joel Asaph Allen]] in 1902.<ref>{{cite journal |last1=Allen |first1=J. A. |year=1902 |title=Mammal names proposed by Oken in his 'Lehrbuch der Zoologie' |journal=Bulletin of the American Museum of Natural History |volume=16 |issue=27 |pages=373−379 |url=http://digitallibrary.amnh.org/items/bc682fbb-d0d7-42a1-9906-bb50b3b0ae31 |access-date=2024-03-14 |archive-date=2023-12-09 |archive-url=https://web.archive.org/web/20231209013613/https://digitallibrary.amnh.org/items/bc682fbb-d0d7-42a1-9906-bb50b3b0ae31 |url-status=live }}</ref>
In 1917, [[Reginald Innes Pocock]] also subordinated the [[tiger]] (''P. tigris''), [[lion]] (''P. leo''), and [[jaguar]] (''P. onca'') to ''Panthera''.<ref>{{cite journal |last1=Pocock |first1=R. I. |year=1917 |title=The Classification of existing Felidae |journal=The Annals and Magazine of Natural History |series=Series 8 |volume=XX |pages=329–350 |doi=10.1080/00222931709487018 |url=https://archive.org/stream/annalsmagazineof8201917lond#page/n359/mode/2up}}</ref><ref>{{cite book |last=Pocock |first= R. I. |year=1939 |title=The Fauna of British India, including Ceylon and Burma. Mammalia: Volume 1 |publisher=Taylor and Francis |___location=London |pages=222–239 |chapter=''Panthera pardus'' |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n273/mode/2up}}</ref>
===Living subspecies===
Following Linnaeus' first description, 27 leopard [[subspecies]] were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered [[Valid name (zoology)|valid]] on the basis of [[mitochondrial]] analysis.<ref name=Miththapala1996>{{cite journal |last1=Miththapala |first1=S. |last2=Seidensticker |first2=J. |last3=O'Brien |first3=S. J. |name-list-style=amp |title=Phylogeographic subspecies recognition in leopards (''Panthera pardus''): molecular genetic variation |journal=Conservation Biology |year=1996 |volume=10 |issue=4 |pages=1115–1132 |doi=10.1046/j.1523-1739.1996.10041115.x |bibcode=1996ConBi..10.1115M |url=https://repository.si.edu/bitstream/handle/10088/4298/Miththapala1996.pdf?sequence=1&isAllowed=y |access-date=2017-03-09 |archive-date=2016-02-17 |archive-url=https://web.archive.org/web/20160217074413/https://repository.si.edu/bitstream/handle/10088/4298/Miththapala1996.pdf?sequence=1&isAllowed=y |url-status=live }}</ref> Later analysis revealed a ninth valid subspecies, the [[Arabian leopard]].<ref name="Uphyrkina">{{cite journal |last1=Uphyrkina |first1=O. |last2=Johnson |first2=E. W. |last3=Quigley |first3=H. |last4=Miquelle |first4=D. |last5=Marker |first5=L. |last6=Bush |first6=M. |last7=O'Brien |first7=S. J. |name-list-style=amp |year=2001 |title=Phylogenetics, genome diversity and origin of modern leopard, ''Panthera pardus'' |url=http://www.biosoil.ru/files/00001386.pdf |url-status=live |journal=[[Molecular Ecology]] |volume=10 |issue=11 |pages=2617–2633 |doi=10.1046/j.0962-1083.2001.01350.x |pmid=11883877 |bibcode=2001MolEc..10.2617U |s2cid=304770 |archive-url=https://web.archive.org/web/20110910162803/http://www.biosoil.ru/files/00001386.pdf |archive-date=2011-09-10}}</ref>
In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:<ref name=CatSG2017>{{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. |name-list-style=amp |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 |issue=Special Issue 11 |pages=73–75 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=73 |access-date=2019-07-25 |archive-date=2020-01-17 |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=73 |url-status=live}}</ref>
{| class="wikitable"
! Subspecies !! Distribution !! Image
|- style="vertical-align: top;"
|[[African leopard]] (''P. p. pardus'') {{small|(Linnaeus, 1758)}}<ref name=msw3/>
||It is the most widespread leopard subspecies and is native to most of [[Sub-Saharan Africa]], but likely [[Local extinction|locally extinct]] in [[Mauritania]], [[Togo]], [[Morocco]], [[Algeria]], [[Tunisia]] and [[Libya]] and most likely also in [[Gambia]] and [[Lesotho]].<ref name=IUCN/>
||[[File:Leopard (Panthera pardus) male ... (51890626416).jpg|frameless]]
|- style="vertical-align: top;"
|[[Indian leopard]] (''P. p. fusca'') {{small|(Meyer, 1794)}}<ref>{{cite book |author=Meyer, F. A. A. |year=1794 |title=Zoologische Annalen. Erster Band |___location=Weimar |publisher=Im Verlage des Industrie-Comptoirs |pages=394–396 |chapter=Über de la Metheries schwarzen Panther |chapter-url=https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394 |access-date=2019-04-05 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145120/https://books.google.com/books?id=v4s-AAAAcAAJ&pg=PA394#v=onepage&q&f=false |url-status=live }}</ref>
||It occurs in the [[Indian subcontinent]], [[Myanmar]] and southern [[Tibet]].<ref name=CatSG2017/><ref>{{cite journal |last1=Laguardia, A. |last2=Kamler, J. F. |last3=Li, S. |last4=Zhang, C. |last5=Zhou, Z. |last6=Shi, K. |year=2017 |title=The current distribution and status of leopards ''Panthera pardus'' in China |journal=Oryx |volume=51 |issue=1 |pages=153−159 |doi=10.1017/S0030605315000988 |doi-access=free |name-list-style=amp}}</ref> It is listed as [[Near-threatened species|Near Threatened]].<ref>{{cite iucn |title=''Panthera pardus ssp. fusca'' |author=Shivakumar, S. |author2=Khettry, A. |author3=Surve, N. |author4=Rahman, H. |author5=Ghimirey, Y. |author6=Tharchen, L. |author7=Zaw, T. |author8=Waseem, M. |author9=Jhala, Y. |name-list-style=amp |date=2023 |page=e.T215195524A215195533 |access-date=17 March 2024}}</ref>
||[[File:Indian male leopard (cropped).jpg|frameless]]
|- style="vertical-align: top;"
|[[Javan leopard]] (''P. p. melas'') {{small|([[Georges Cuvier|Cuvier]], 1809)}}<ref>{{cite journal |author=Cuvier, G. |year=1809 |title=Recherches sur les espėces vivantes de grands chats, pour servir de preuves et d'éclaircissement au chapitre sur les carnassiers fossils |journal=Annales du Muséum National d'Histoire Naturelle |volume=Tome XIV |pages=136–164}}</ref>
||It is native to [[Java]] in [[Indonesia]] and has been assessed as [[Endangered species|Endangered]] in 2021.<ref>{{cite iucn |title=''Panthera pardus'' ssp. ''melas'' |name-list-style=amp |author=Wibisono, H. |author2=Wilianto, E. |author3=Pinondang, I. |author4=Rahman, D.A. |author5=Chandradewi, D. |year=2021 |page=e.T15962A50660931 |doi=10.2305/IUCN.UK.2021-2.RLTS.T15962A50660931.en |access-date=11 November 2023}}</ref>
||[[File:IG KusumoKintokoEko WA 082140100111 foto macan tutul jawa lokasi TN Baluran, Situbondo, Indonesia.jpg|frameless]]
|- style="vertical-align: top;"
|[[Arabian leopard]] (''P. p. nimr'') {{small|([[Wilhelm Hemprich|Hemprich]] and [[Christian Gottfried Ehrenberg|Ehrenberg]], 1830)}}<ref>{{cite book |last1=Hemprich |first1=W. |last2=Ehrenberg |first2=C. G. |year=1830 |chapter=Felis, pardus?, nimr |pages=Plate 17 |title=Symbolae Physicae, seu Icones et Descriptiones Mammalium quae ex Itinere per Africam Borealem et Asiam Occidentalem Friderici Guilelmi Hemprich et Christiani Godofredi Ehrenberg. Decas Secunda. Zoologica I. Mammalia II |___location=Berolini |publisher=Officina Academica |editor=Dr. C. G. Ehrenberg |chapter-url=https://archive.org/details/SymbolaephysicaMammEhreA/page/n60}}</ref>
||It is the smallest leopard subspecies and considered [[Endemism|endemic]] to the [[Arabian Peninsula]].<ref>{{cite journal |last1=Spalton, J. A. |last2=Al Hikmani, H. M. |name-list-style=amp |year=2006 |title=The Leopard in the Arabian Peninsula – Distribution and subspecies status |journal=Cat News |issue=Special Issue 1 |pages=4–8 |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.2._Status_Reports/leopard/Spalton___Hikmani_2006_Status_of_the_Leopard_on_the_Arabian_Peninsula.pdf |archive-url=https://web.archive.org/web/20150619075145/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.2._Status_Reports/leopard/Spalton___Hikmani_2006_Status_of_the_Leopard_on_the_Arabian_Peninsula.pdf |archive-date=2015-06-19 |url-status=live}}</ref> As of 2023, the population was estimated to comprise 100–120 individuals in [[Oman]] and [[Yemen]]; it was therefore assessed as [[Critically Endangered]] in 2023.<ref>{{cite iucn |title=''Panthera pardus ssp. nimr'' |name-list-style=amp |author=Al Hikmani, H. |author2=Spalton, A. |author3=Zafar-ul Islam, M. |author4=al-Johany, A. |author5=Sulayem, M. |author6=Al-Duais, M. |author7=Almalki, A. |date=2023 |page=e.T15958A46767457 |access-date=17 March 2024}}</ref> It is locally extinct in Syria, Lebanon, Israel, Palestine, Jordan, Kuwait and the United Arab Emirates.<ref name=IUCN/>
||[[File:Arabian Leopard Panthera pardus nimr in Sharjah Photo Prof Dr Norman Ali Khalaf.jpg|frameless]]
|- style="vertical-align: top;"
|[[Panthera pardus tulliana|''P. p. tulliana'']] {{small|([[Achille Valenciennes|Valenciennes]], 1856)}}<ref name=Valenciennes>{{cite journal |author=Valenciennes, A. |date=1856 |title=Sur une nouvelles espèce de Panthère tué par M. Tchihatcheff à Ninfi, village situé à huit lieues est de Smyrne |journal=Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences |volume=42 |pages=1035–1039}}</ref>
||It occurs from eastern [[Turkey]] and the [[Caucasus]] to the [[Iranian Plateau]] and the [[Hindu Kush]] into the western [[Himalayas]]. It is listed as [[Endangered]].<ref>{{cite iucn |title=''Panthera pardus ssp. tulliana'' |name-list-style=amp |author=Ghoddousi, A. |author2=Khorozyan, I. |date=2023 |page=e.T15961A50660903 |doi=10.2305/IUCN.UK.2023-1.RLTS.T15961A50660903.en |access-date=17 March 2024}}</ref> It is locally extinct in [[Uzbekistan]] and [[Tajikistan]].<ref name=IUCN/>
The Balochistan leopard population in the south of Iran, Afghanistan and Pakistan is separated from the northern population by the [[Dasht-e Kavir]] and [[Dasht-e Lut]] deserts.<ref name=Khorozyan2006>{{cite journal |last=Khorozyan |first=I. G. |author2=Gennady, F. |author3=Baryshnikov, G. F. |author4=Abramov, A. V. |name-list-style=amp |year=2006 |title=Taxonomic status of the leopard, ''Panthera pardus'' (Carnivora, Felidae) in the Caucasus and adjacent areas |journal=Russian Journal of Theriology |volume=5 |issue=1 |pages=41–52 |doi=10.15298/rusjtheriol.05.1.06 |doi-access=free}}</ref>
||<span style="{{MirrorH}}">[[File:Nordpersischen Leoparden.jpg|frameless]]</span>
|- style="vertical-align: top;"
|[[Amur leopard]] (''P. p. orientalis'') {{small|([[Hermann Schlegel|Schlegel]], 1857)}}<ref>{{cite book |author=Schlegel, H. |year=1857 |chapter=''Felis orientalis'' |page=23 |title=Handleiding Tot de Beoefening der Dierkunde, I<sup>e</sup> Deel |publisher=Boekdrukkerij van Nys |___location=Breda}}</ref><ref>{{cite journal |author=Gray, J. E. |year=1862 |title=Description of some new species of Mammalia |journal=Proceedings of the Royal Zoological Society of London |volume=30 |pages=261−263, plate XXXIII |doi=10.1111/j.1469-7998.1862.tb06524.x |url=https://archive.org/details/proceedingsofgen62zool/page/262}}</ref>
||It is native to the [[Russian Far East]] and [[northern China]], but is locally extinct in the [[Korean peninsula]].<ref name=IUCN/>
||<span style="{{MirrorH}}">[[File:Amur leopard. Frame from a camera trap (cropped).jpg|frameless]]</span>
|- style="vertical-align: top;"
|[[Indochinese leopard]] (''P. p. delacouri'') {{small|[[Reginald Innes Pocock|Pocock]], 1930}}<ref>{{cite journal |author=Pocock, R. I. |year=1930 |title=The Panthers and Ounces of Asia |journal=Journal of the Bombay Natural History Society |volume=34 |issue=2 |pages=307–336}}</ref>
||It occurs in mainland Southeast Asia and [[southern China]], and is listed as [[Critically Endangered]].<ref>{{cite iucn |title=''Panthera pardus'' ssp. ''delacouri'' |author1=Rostro-García, S. |author2=Kamler, J.F. |author3=Clements, G.R. |author4=Lynam, A.J. |author5=Naing, H. |name-list-style=amp |year=2019 |page=e.T124159083A163986056 |access-date=17 March 2024}}</ref> It is locally extinct in [[Hong Kong]], [[Singapore]], [[Laos]] and [[Vietnam]].<ref name=IUCN/>
||[[File:Indochinese leopard.jpg|frameless]]
|- style="vertical-align: top;"
|[[Sri Lankan leopard]] (''P. p. kotiya'') {{small|[[Paulus Edward Pieris Deraniyagala|Deraniyagala]], 1956}}<ref>{{cite journal |author=Deraniyagala, P. E. P. |year=1956 |title=The Ceylon leopard, a distinct subspecies |journal=Spolia Zeylanica |volume=28 |pages=115–116}}</ref>
||It is native to [[Sri Lanka]] and listed as [[Vulnerable species|Vulnerable]].<ref>{{cite iucn |title=''Panthera pardus ssp.kotiya'' |name-list-style=amp |author=Kittle, A.M. |author2=Watson, A. |date=2020 |page=e.T15959A50660847 |access-date=17 March 2024}}</ref>
||[[File:Srilankan leopard (srilankan kotiya) 02 (cropped).jpg|frameless]]
|}
Results of an [[analysis of molecular variance]] and pairwise [[fixation index]] of 182 African leopard museum specimens showed that some African leopards exhibit higher genetic differences than Asian leopard subspecies.<ref>{{cite journal |author1=Anco, C. |author2=Kolokotronis, S. O. |author3=Henschel, P. |author4=Cunningham, S. W. |author5=Amato, G. |author6=Hekkala, E. |name-list-style=amp |title=Historical mitochondrial diversity in African leopards (''Panthera pardus'') revealed by archival museum specimens |journal=Mitochondrial DNA Part A |volume=29 |issue=3 |pages=455–473 |doi=10.1080/24701394.2017.1307973 |pmid=28423965 |year=2017 |s2cid=4348541}}</ref>
===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 |bibcode=2010NJGPA.258..339M |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 |last1=Mecozzi |first1=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 |article-number=106840 |doi=10.1016/j.quascirev.2021.106840 |bibcode=2021QSRv..26206840M |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 |article-number=e12020 |doi=10.7717/peerj.12020 |pmid=34513335 |pmc=8388558 |doi-access=free}}</ref>
===Hybrids===
{{Main|Panthera hybrid|Pumapard}}
In 1953, a male leopard and a female lion were [[Crossbreeding|crossbred]] in Hanshin Park in [[Nishinomiya, Japan]]. Their offspring known as a [[leopon]] was born in 1959 and 1961, all cubs were spotted and bigger than a juvenile leopard. Attempts to mate a leopon with a tigress proved unsuccessful.<ref>{{cite book |editor1=Kisling, V.N. |title=Zoo and Aquarium History : Ancient Animal Collections to Zoological Gardens |year=2001 |publisher=CRC Press |___location=Boca Raton, Florida |isbn=978-0-8493-2100-9 |author=Kawata, K. |chapter=Zoological gardens of Japan |pages=295–329 |chapter-url=https://books.google.com/books?id=ulbMBQAAQBAJ&pg=PA314 |access-date=2021-09-07 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145322/https://books.google.com/books?id=ulbMBQAAQBAJ&pg=PA314#v=onepage&q&f=false |url-status=live }}</ref>
==Characteristics==
{{Multiple image |direction=vertical |align=right|image1=Leopard skeleton (black background).jpg|caption1=Mounted skeleton |image2=Panthera pardus 3d scan Natural History Museum University of Pisa C 1389.stl|caption2=3D model of skeleton|image3=Persian Leopard Fur 02.JPG |caption3=Rosettes of a leopard|image4=Blackleopard.JPG|caption4=A melanistic leopard or black panther}}
The leopard's fur is generally soft and thick, notably softer on the belly than on the back.<ref name=Skinner>{{cite book |year=2005 |title=The mammals of the southern African sub region |___location=Cambridge |publisher=Cambridge University Press |isbn=9780521844185 |editor1-last=Skinner |editor1-first=J. D. |editor2-last=Chimimba |editor2-first=C. T. |edition=Third |last1=Mills |first1=M. G. L. |chapter=Subfamily Pantherinae |pages=385–396 |chapter-url=https://books.google.com/books?id=iqwEYkTDZf4C&pg=PA385 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150034/https://books.google.com/books?id=iqwEYkTDZf4C&pg=PA385#v=onepage&q&f=false |url-status=live}}</ref> Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in [[Rosette (zoology)|rosette]]s. Its underbelly is white and its ringed tail is shorter than its body. Its pupils are round.<ref name=Mivart1900>{{cite book |last1=Mivart |first1=St. G. J. |author-link=St. George Jackson Mivart |title=The Cat: An Introduction to the Study of Backboned Animals, Especially Mammals |date=1900 |publisher=John Murray |___location=London |chapter=Different kind of Cats |pages=391–439 |chapter-url=https://archive.org/details/catintroductiont00miva/page/394}}</ref> Leopards living in arid regions are pale cream, yellowish to [[ochraceous]] and [[rufous]] in colour; those living in forests and mountains are much darker and deep golden. Spots fade toward the white underbelly and the insides and lower parts of the legs.<ref name=Pocock1932>{{cite journal |last1=Pocook |first1=R. I. |year=1932 |title=The Leopards of Africa |journal=Proceedings of the Zoological Society of London |volume=102 |issue=2 |pages=543–591 |doi=10.1111/j.1096-3642.1932.tb01085.x}}</ref> Rosettes are circular in East African leopard populations, and tend to be squarish in Southern African and larger in Asian leopard populations. The fur tends to be grayish in colder climates, and dark golden in [[rainforest]] habitats.<ref name=CAP>{{cite book |author=Nowell, K. |author2=Jackson, P. |name-list-style=amp |year=1996 |title=Wild Cats: status survey and conservation action plan |publisher=IUCN/SSC Cat Specialist Group |___location=Gland, Switzerland |chapter=Leopard ''Panthera pardus'' (Linnaeus, 1758) |archive-url=https://web.archive.org/web/20140222223654/http://lynx.uio.no/lynx/catsgportal/cat-website/catfolk/ssaprd01.htm |archive-date=2014-02-22 |chapter-url=http://lynx.uio.no/lynx/catsgportal/cat-website/catfolk/ssaprd01.htm}}</ref> Rosette patterns are unique in each individual.<ref name=Schutze>{{cite book |last1=Schütze |first1=H. |title=Field Guide to the Mammals of the Kruger National Park |date=2002 |publisher=Struik Publishers|___location=Cape Town, South Africa |isbn=978-1-86872-594-6 |pages=92–93}}</ref><ref>{{cite book|last1=Menon |first1=V. |title=Indian Mammals: A Field Guide |date=2014 |publisher=Hachette |___location=Gurgaon, India |isbn=978-93-5009-761-8}}</ref> This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as [[camouflage]].<ref>{{cite journal |author1=Allen, W. L. |author2=Cuthill, I. C. |author3=Scott-Samuel, N. E. |author4=Baddeley, R. |year=2010 |name-list-style=amp |title=Why the leopard got its spots: relating pattern development to ecology in felids |journal=Proceedings of the Royal Society B |volume=278 |issue=1710 |pages=1373–1380 |doi=10.1098/rspb.2010.1734 |pmc=3061134 |pmid=20961899}}</ref>
Its white-tipped tail is about {{cvt|60|-|100|cm|1}} long, white underneath and with spots that form incomplete bands toward the end of the tail.<ref name=Hoath>{{cite book |last1=Hoath |first1=R. |title=Field Guide to the Mammals of Egypt |date=2009 |publisher=American University in Cairo Press |___location=Cairo, Egypt |isbn=978-977-416-254-1 |pages=106–107 |chapter=Leopard ''Panthera pardus'' (Linnaeus, 1758) |chapter-url=https://books.google.com/books?id=agWfg6oEKKkC&pg=PA106 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145927/https://books.google.com/books?id=agWfg6oEKKkC&pg=PA106#v=onepage&q&f=false |url-status=live}}</ref>
The guard hairs protecting the basal hairs are short, {{cvt|3-4|mm|1}} in face and head, and increase in length toward the flanks and the belly to about {{cvt|25|-|30|mm|1}}. Juveniles have woolly fur that appear to be dark-coloured due to the densely arranged spots.<ref name=Schutze/><ref name=estes>{{cite book |last=Estes |first=R. |year=1991 |title=The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates |___location=Los Angeles |publisher=The University of California Press |isbn=978-0-520-08085-0 |chapter=Leopard ''Panthera pardus'' |pages=366–369 |chapter-url=https://books.google.com/books?id=g977LsZHpcsC&pg=PA366 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145853/https://books.google.com/books?id=g977LsZHpcsC&pg=PA366#v=onepage&q&f=false |url-status=live}}</ref>
Its fur tends to grow longer in colder climates.<ref name=mammal>{{cite journal |author1=Stein, A. B. |name-list-style=amp |author2=Hayssen, V. |title=''Panthera pardus'' (Carnivora: Felidae) |journal=[[Mammalian Species]] |date=2010 |volume=45 |issue=900 |pages=30–48 |doi=10.1644/900.1 |s2cid=44839740 |doi-access=free|pmc=7149468 }}</ref>
The leopard's rosettes differ from those of the [[jaguar]], which are darker and with smaller spots inside.<ref name=Mivart1900/> The leopard has a [[Diploidy|diploid]] [[chromosome]] number of 38.<ref name=HeptnerSludskiy1972>{{Cite book |last1=Heptner |first1=V. G. |name-list-style=amp |last2=Sludskii |first2=A. A. |orig-year=1972 |year=1992 |title=Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola |trans-title=Mammals of the Soviet Union, Volume II, Part 2 |publisher=Smithsonian Institution and the National Science Foundation |___location=Washington DC |chapter=Bars (leopard) |chapter-url=https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up |pages=203–273 |isbn=978-90-04-08876-4}}</ref>
Melanistic leopards are also known as [[black panther]]s. [[Melanism]] in leopards is caused by a [[recessive allele]] and is inherited as a [[recessive trait]].<ref>{{cite journal|last1=Robinson |first1=R. |title=Inheritance of the black form of the leopard ''Panthera pardus'' |journal=Genetica |year=1970 |volume=41 |issue=1|pages=190–197 |doi=10.1007/BF00958904 |pmid=5480762 |s2cid=5446868}}</ref><ref name=Eizirik>{{cite journal |last1=Eizirik |first1=E. |last2=Yuhki |first2=N. |last3=Johnson |first3=W. E.|last4=Menotti-Raymond |first4=M. |last5=Hannah |first5=S. S. |last6=O'Brien |first6=S. J. |title=Molecular genetics and evolution of melanism in the cat family |journal=Current Biology |year=2003 |volume=13 |issue=5 |pages=448–453 |doi=10.1016/S0960-9822(03)00128-3 |pmid=12620197 |s2cid=19021807 |doi-access=free |bibcode=2003CBio...13..448E}}</ref><ref>{{cite journal |last1=Kawanishi |first1=K. |last2=Sunquist |first2=M. E. |last3=Eizirik |first3=E. |last4=Lynam |first4=A. J. |last5=Ngoprasert |first5=D. |last6=Wan Shahruddin |first6=W. N. |last7=Rayan |first7=D. M. |last8=Sharma |first8=D. S. K.|last9=Steinmetz|first9=R. |name-list-style=amp |title=Near fixation of melanism in leopards of the Malay Peninsula |journal=Journal of Zoology |year=2010 |volume=282 |issue=3 |pages=201–206 |doi=10.1111/j.1469-7998.2010.00731.x}}</ref><ref name=daSilva17>{{cite journal |author1=Da Silva L. G. |author2=Kawanishi, K. |first=K. |author3=Henschel P. |author4=Kittle, A. |author5=Sanei, A. |author6=Reebin, A. |author7=Miquelle, D. |author8=Stein, A. B. |author9=Watson, A. |author10=Kekule, L. B. |author11=Machado, R. B. |author12=Eizirik, E. |name-list-style=amp |year=2017 |title=Mapping black panthers: Macroecological modeling of melanism in leopards (''Panthera pardus'') |journal= PLOS ONE |volume=12 |issue=4 |pages=e0170378 |doi=10.1371/journal.pone.0170378 |pmid=28379961 |pmc=5381760 |bibcode=2017PLoSO..1270378D |doi-access=free}}</ref>
In India, nine pale and white leopards were reported between 1905 and 1967.<ref>{{cite journal |author=Divyabhanusinh |year=1993 |title=On mutant leopards ''Panthera pardus'' from India |journal=Journal of the Bombay Natural History Society |volume=90 |issue=1 |pages=88−89 |url=https://archive.org/details/journalofbombay901993bomb/page/88}}</ref>
Leopards exhibiting [[erythrism]] were recorded between 1990 and 2015 in South Africa's [[Madikwe Game Reserve]] and in [[Mpumalanga]]. The cause of this [[Polymorphism (biology)|morph]] known as a "strawberry leopard" or "pink panther" is not well understood.<ref>{{Cite journal |last1=Pirie |first1=T. J. |last2=Thomas |first2=R. L. |last3=Fellowes |first3=M. D. E. |name-list-style=amp |date=2016 |title=Erythristic leopards ''Panthera pardus'' in South Africa |journal=Bothalia |volume=46 |issue=1 |pages=1–5 |doi=10.4102/abc.v46i1.2034 |doi-access=free}}</ref>
===Size===
The leopard is a slender and muscular cat, with relatively short limbs and a broad head. It is [[sexually dimorphic]] with males larger and heavier than females.<ref name=Hoath/> Males stand {{cvt|60|-|70|cm}} at the shoulder, while females are {{cvt|57|-|64|cm}} tall. The head-and-body length ranges between {{cvt|92|and|183|cm}} with a {{cvt|66|to|102|cm|}} long tail. Sizes vary geographically. Males typically weigh {{cvt|30.9|-|72|kg}}, and females {{cvt|20.5|-|43|kg}}.<ref name="Kingdon">{{cite book |author1=Hunter, L. |author2=Henschel, P. |author3=Ray, J. C. |name-list-style=amp |chapter=''Panthera pardus'' Leopard |pages=159–168 |date=2013 |isbn=978-1-4081-8996-2 |chapter-url=https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA159 |editor1=Kingdon, J. |editor2=Happold, D. |editor3=Butynski, T. |editor4=Hoffmann, M. |editor5=Happold, M. |editor6=Kalina, J. |title=Mammals of Africa |publisher=Bloomsbury Publishing |___location=London |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145933/https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA159#v=onepage&q&f=false |url-status=live}}</ref> Occasionally, large males can grow up to {{cvt|91|kg}}. Leopards from the [[Cape Province]] in South Africa are generally smaller, reaching only {{cvt|20|-|45|kg}} in males.<ref name=estes/><ref name=mammal /><ref name=nowak>{{cite book |last1=Nowak |first1=R. M. |title=Walker's Mammals of the World |year=1999 |publisher=Johns Hopkins University Press |___location=Baltimore, US |isbn=978-0-8018-5789-8 |edition=Sixth |pages=828–831 |chapter=''Panthera pardus'' (Leopard) |chapter-url=https://books.google.com/books?id=T37sFCl43E8C&pg=PA828 |access-date=2021-05-19 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145930/https://books.google.com/books?id=T37sFCl43E8C&pg=PA828#v=onepage&q&f=false |url-status=live}}</ref>
The heaviest wild leopard in Southern Africa weighed around {{cvt|96|kg}}, and it measured {{cvt|262|cm}}.<ref name=Burnie>{{cite book |editor-last1=Burnie |editor-first1=D. |editor-last2=Wilson |editor-first2=D. E. |name-list-style=amp |year=2001 |title=Animal: The Definitive Visual Guide to the World's Wildlife |publisher=DK Adult |isbn=978-0-7894-7764-4}}</ref> In 2016, an Indian leopard killed in [[Himachal Pradesh]] measured {{cvt|261|cm}} with an estimated weight of {{cvt|78.5|kg}}; it was perhaps the largest known wild leopard in India.<ref>{{cite news |year=2016 |title=Is this the longest leopard in India? |newspaper=The Times of India |url=https://timesofindia.indiatimes.com/city/chandigarh/Is-this-the-longest-leopard-in-India/articleshow/56227308.cms |access-date=2018-03-14 |archive-date=2019-04-06 |archive-url=https://web.archive.org/web/20190406154410/https://timesofindia.indiatimes.com/city/chandigarh/Is-this-the-longest-leopard-in-India/articleshow/56227308.cms |url-status=live}}</ref>
The largest recorded skull of a leopard was found in India in 1920 and measured {{cvt|28|cm}} in [[Basal (anatomy)|basal]] length, {{cvt|20|cm}} in breadth, and weighed {{cvt|1|kg}}. The skull of an African leopard measured {{cvt|286|mm}} in basal length, and {{cvt|181|mm}} in breadth, and weighed {{cvt|790|g}}.<ref>{{cite journal |author=Prater, S. H. |title=Record panther skull (''P. p. pardus'') |journal=The Journal of the Bombay Natural History Society |date=1921 |volume=XXVII |issue=Part IV |pages=933–935 |url=https://www.biodiversitylibrary.org/item/95740#page/1055/mode/1up |access-date=2019-08-24 |archive-date=2019-05-02 |archive-url=https://web.archive.org/web/20190502064501/https://www.biodiversitylibrary.org/item/95740#page/1055/mode/1up |url-status=live}}</ref>
== Distribution and habitat ==
[[File:David Raju Leopard0827.jpg|thumb|Leopard in a tree in India]]
[[File:Edgar Grooming (49611421486).jpg|thumb|An [[Amur leopard]] at the [[Cleveland Metroparks Zoo]] [[personal grooming|grooming himself]]]]
The leopard has the largest distribution of all wild cats, occurring widely in Africa and Asia, although populations are fragmented and declining.<ref name=IUCN/> It inhabits foremost [[savanna]] and [[rainforest]], and areas where [[grassland]]s, [[woodland]]s and [[riparian forest]]s remain largely undisturbed.<ref name=CAP/> It also persists in urban environments, if it is not persecuted, has sufficient prey and patches of vegetation for shelter during the day.<ref name=Powell2021>{{Cite journal |last1=Powell |first1=J. |last2=Axmacher |first2=J. C. |last3=Linnell |first3=J. D.C. |last4=Durant |first4=S. M. |title=Diverse locations and a long history: Historical context for urban Leopards (''Panthera pardus'') in the early anthropocene From Seoul, Korea |year=2021 |doi=10.3389/fcosc.2021.765911 |journal=Frontiers in Conservation Science |volume=2 |article-number=765911 |bibcode=2021FrCS....2.5911P |doi-access=free |name-list-style=amp |hdl=11250/2999764 |hdl-access=free}}</ref>
The leopard's range in [[West Africa]] is estimated to have drastically declined by 95%, and in the [[Sahara desert]] by 97%.<ref name=Jacobson2016>{{cite journal |last1=Jacobson|first1=A. P. |last2=Gerngross |first2=P. |last3=Lemeris |first3=J. R. Jr.|last4=Schoonover |first4=R. F. |name-list-style=amp |last5=Anco |first5=C. |last6=Breitenmoser-Würsten |first6=C. |last7=Durant |first7=S. M. |last8=Farhadinia |first8=M. S. |last9=Henschel |first9=P. |last10=Kamler |first10=J. F. |last11=Laguardia |first11=A. |last12=Rostro-García |first12=S. |last13=Stein |first13=A. B. |last14=Dollar |first14=L. |title=Leopard (''Panthera pardus'') status, distribution, and the research efforts across its range |journal=PeerJ |date=2016 |volume=4 |article-number=e1974 |doi=10.7717/peerj.1974 |pmid=27168983 |pmc=4861552 |doi-access=free}}</ref> In [[sub-Saharan Africa]], it is still numerous and surviving in marginal habitats where other large cats have disappeared.<ref>{{Cite journal |last1=Pirie |first1=T. J. |last2=Thomas |first2=R. L. |last3=Fellowes |first3=M. D. E. |name-list-style=amp |date=2017 |title=Increasing game prices may alter farmers' behaviours towards leopards (''Panthera pardus'') and other carnivores in South Africa |article-number=e3369 |journal=PeerJ |volume=5 |doi=10.7717/peerj.3369|pmid=28584709 |pmc=5452990 |doi-access=free}}</ref> In southeastern [[Egypt]], an individual found killed in 2017 was the first sighting of the leopard in this area in 65 years.<ref>{{Cite journal |last1=Soultan, A. |last2=Attum, O. |last3=Hamada, A. |last4=Hatab, E. B. |last5=Ahmed, S. E. |last6=Eisa, A. |last7=Al Sharif, I. |last8=Nagy, A. |name-list-style=amp |author9=Shohdi, W. |date=2017 |title=Recent observation for leopard ''Panthera pardus'' in Egypt |journal=Mammalia |volume=81 |issue=1 |pages=115–117 |doi=10.1515/mammalia-2015-0089 |s2cid=90676105}}</ref>
In [[West Asia]], the leopard inhabits the areas of southern and southeastern [[Anatolia]].<ref>{{cite journal |author1=Toyran, K. |title=Noteworthy record of ''Panthera pardus'' in Turkey (Carnivora: Felidae) |journal=Fresenius Environmental Bulletin |year=2018 |volume=27 |issue=11 |pages=7348–7353}}</ref><ref>{{cite journal |author1=Karataş, A. |name-list-style=amp |author2=Bulut, Ş. |author3=Akbaba, B. |year=2021 |title=Camera trap records confirm the survival of the Leopard (''Panthera pardus'' L., 1758) in eastern Turkey (Mammalia: Felidae) |journal=Zoology in the Middle East |volume= 67 |issue= 3|pages=198–205 |s2cid=235564429 |doi=10.1080/09397140.2021.1924419}}</ref><ref>{{Cite journal |title=New records of the living Anatolian Leopard (''Panthera pardus tulliana'' L., 1758) in the Mediterranean region of Turkey |author1=Ünal, Y. |author2=Uysal, H. |author3=Koca, A. |author4=Zenbilci, M. |journal=Applied Ecology and Environmental Research |volume=21 |pages=1043–1059 |issue=2 |doi=10.15666/aeer/2102_10431059 |date=2023 |bibcode=2023ApEER..21.1043U |doi-access=free |name-list-style=amp}}</ref>
Leopard populations in the [[Arabian Peninsula]] are small and fragmented.<ref>{{cite journal |author1=Spalton, J. A. |author2=Al Hikmani, H. M. |name-list-style=amp |year=2006 |title=The Leopard in the Arabian Peninsula – Distribution and subspecies status |journal=Cat News |issue=Special Issue 1 |pages=4–8 |url=http://catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.1._SI_1/Spalton___Al_Hikmani_2006_Leopard_in_the_Arabian_Peninsula.pdf }}</ref><ref>{{cite journal |last1=Judas |first1=J. |last2=Paillat |first2=P. |last3=Khoja |first3=A. |last4=Boug |first4=A. |name-list-style=amp |year=2006 |title=Status of the Arabian leopard in Saudi Arabia |journal=Cat News |issue=Special Issue 1 |pages=11–19 |url=http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.2._Status_Reports/leopard/Judas_et_al_2006_Status_of_the_Arabian_Leopard_in_Saudi_Arabia.pdf |archive-date=2015-09-19 |access-date=2019-03-17 |archive-url=https://web.archive.org/web/20150919114352/http://www.catsg.org/fileadmin/filesharing/3.Conservation_Center/3.2._Status_Reports/leopard/Judas_et_al_2006_Status_of_the_Arabian_Leopard_in_Saudi_Arabia.pdf |url-status=dead }}</ref><ref name=AlJ06>{{cite journal |last1=Al Jumaily, M. |last2=Mallon, D. P. |last3=Nasher, A. K. |last4=Thowabeh, N. |name-list-style=amp |year=2006 |title=Status Report on Arabian Leopard in Yemen |journal=Cat News |issue=Special Issue 1 |pages=20–25 |url=http://catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.1._SI_1/Jumaily_et_al_2006_Status_Report_on_Arabian_Leopard_in_Yemen.pdf}}</ref>
In the [[Indian subcontinent]], the leopard is still relatively abundant, with greater numbers than those of other ''Panthera'' species.<ref name=IUCN/> Some leopard populations in [[India]] live quite close to human settlements and even in semi-developed areas. Although adaptable to human disturbances, leopards require healthy prey populations and appropriate vegetative cover for hunting for prolonged survival and thus rarely linger in heavily developed areas. Due to the leopard's stealth, people often remain unaware that it lives in nearby areas.<ref name=Arthreya>{{cite web |author=Arthreya, V. |year=2012 |title=Living with Leopards Outside Protected Areas in India |url=http://www.conservationindia.org/articles/living-with-leopards-outside-protected-areas-in-india |publisher=Conservation India |access-date=2012-12-10 |archive-date=2012-12-21 |archive-url=https://web.archive.org/web/20121221051725/http://www.conservationindia.org/articles/living-with-leopards-outside-protected-areas-in-india |url-status=live}}</ref> As of 2020, the leopard population within forested habitats in India's tiger range landscapes was estimated at 12,172 to 13,535 individuals. Surveyed landscapes included elevations below {{cvt|2600|m}} in the [[Shivalik Hills]] and [[Gangetic plains]], [[Central India]] and [[Eastern Ghats]], [[Western Ghats]], the [[Brahmaputra River]] basin and hills in [[Northeast India]].<ref>{{cite report |author=Jhala, Y.V. |author-link=Yadvendradev Vikramsinh Jhala |author2=Qureshi, Q. |author3=Yadav, S.P. |name-list-style=amp |year=2020 |title=Status of leopards in India, 2018. Technical Report TR/2020/16 |___location=New Delhi and Dehradun |publisher=National Tiger Conservation Authority, Government of India and Wildlife Institute of India}}</ref>
In Nepal's [[Kanchenjunga Conservation Area]], a melanistic leopard was photographed at an elevation of {{cvt|4300|m}} by a camera trap in May 2012.<ref>{{cite journal |author1=Thapa, K. |author2=Pradhan, N. M. B. |author3=Berker, J. |author4=Dhakal, M. |author5=Bhandari, A. R. |author6=Gurung, G. S. |author7=Rai, D. P. |author8=Thapa, G. J. |author9=Shrestha, S. |author10=Singh, G. R. |name-list-style=amp |year=2013 |title=High elevation record of a leopard cat in the Kangchenjunga Conservation Area, Nepal |journal=Cat News |issue=58 |pages=26–27}}</ref>
In Sri Lanka, leopards were recorded in [[Yala National Park]] and in unprotected forest patches, [[tea estate]]s, grasslands, home gardens, [[Pinus|pine]] and [[eucalyptus]] plantations.<ref>{{cite journal |author=Kittle, A. M. |author2=Watson, A. C. |author3=Chanaka Kumara, P. H. |author4=Nimalka Sanjeewani, H. K. |name-list-style=amp |year=2014 |title=Status and distribution of the leopard in the central hills of Sri Lanka |journal=Cat News |issue=56 |pages=28−31}}</ref><ref name=Kittle_al2014>{{cite journal |last1=Kittle |first1=A. M. |last2=Watson |first2=A. C. |last3=Kumara |first3=P. H. S. C. |last4=Sandanayake |first4=S. D. K. C. |last5=Sanjeewani |first5=H. K. N. |last6=Fernando |first6=T. S. P. |name-list-style=amp |year=2014 |title=Notes on the diet and habitat selection of the Sri Lankan Leopard ''Panthera pardus kotiya'' (Mammalia: Felidae) in the central highlands of Sri Lanka |journal=Journal of Threatened Taxa |volume=6 |issue=9 |pages=6214–6221 |doi=10.11609/JoTT.o3731.6214-21 |doi-access=free}}</ref>
In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's [[Karen State]].<ref>{{cite journal |author1=Saw Sha Bwe Moo |author2=Froese, G.Z.L. |author3=Gray, T.N.E. |name-list-style=amp |year=2017 |title=First structured camera-trap surveys in Karen State, Myanmar, reveal high diversity of globally threatened mammals |journal=Oryx |volume=52 |issue=3 |pages=537−543 |doi=10.1017/S0030605316001113 |doi-access=free}}</ref> The Northern [[Tenasserim Hills|Tenasserim]] Forest Complex in southern Myanmar is considered a leopard stronghold. In Thailand, leopards are present in the [[Western Forest Complex]], [[Kaeng Krachan National Park|Kaeng Krachan]]-[[Kui Buri National Park|Kui Buri]], [[Khlong Saeng Wildlife Sanctuary|Khlong Saeng]]-[[Khao Sok National Park|Khao Sok]] protected area complexes and in [[Hala Bala Wildlife Sanctuary]] bordering Malaysia. In [[Peninsular Malaysia]], leopards are present in [[Belum-Temengor]], [[Taman Negara National Park|Taman Negara]] and [[Endau-Rompin National Park]]s.<ref>{{cite journal | last1=Rostro-García |first1=S. |last2=Kamler |first2 =J. F. |last3=Ash |first3=E. |last4=Clements |first4=G. R. |last5=Gibson |first5=L. |last6=Lynam |first6=A. J. |last7=McEwin |first7=R. |last8=Naing |first8=H. |last9=Paglia |first9=S. |name-list-style=amp |year=2016 |title=Endangered leopards: Range collapse of the Indochinese leopard (''Panthera pardus delacouri'') in Southeast Asia |journal=Biological Conservation |volume=201 |pages=293–300 |doi=10.1016/j.biocon.2016.07.001|bibcode=2016BCons.201..293R |hdl=10722/232870 |hdl-access=free}}</ref>
In Laos, leopards were recorded in [[Nam Et-Phou Louey]] National Biodiversity Conservation Area and Nam Kan National Protected Area.<ref>{{cite journal |author=Johnson, A. |author2=Vongkhamheng, C. |name-list-style=amp |author3=Hedemark, M. |author4=Saithongdam, T. |year=2006 |title=Effects of human–carnivore conflict on tiger (''Panthera tigris'') and prey populations in Lao PDR |journal=Animal Conservation |volume=9 |issue=4 |pages=421–430 |doi=10.1111/j.1469-1795.2006.00049.x |bibcode=2006AnCon...9..421J |s2cid=73637721 |url=http://faculty.nelson.wisc.edu/treves/pubs/Arlyne/animal_cons.pdf |archive-url=https://web.archive.org/web/20170810101758/http://faculty.nelson.wisc.edu/treves/pubs/Arlyne/animal_cons.pdf |archive-date=2017-08-10 |url-status=live}}</ref><ref>{{cite book |author=Robichaud, W. |name-list-style=amp |author2=Insua-Cao |author3=Sisomphane, P. C. |author4=Chounnavanh, S. |year=2010 |title=A scoping mission to Nam Kan National Protected Area, Lao PDR |publisher=Fauna & Flora International |chapter=Appendix 4 |pages=33−42 |chapter-url=https://studylib.net/doc/18697244/a-scoping-mission-to-nam-kan-national-protected-area--lao... |access-date=2019-04-07 |archive-date=2019-04-07 |archive-url=https://web.archive.org/web/20190407154414/https://studylib.net/doc/18697244/a-scoping-mission-to-nam-kan-national-protected-area--lao... |url-status=live}}</ref>
In Cambodia, leopards inhabit deciduous [[dipterocarp forest]] in [[Phnom Prich Wildlife Sanctuary]] and [[Mondulkiri Protected Forest]].<ref>{{cite journal |last1=Gray |first1=T. N. |last2=Phan |first2=C. |name-list-style=amp |year=2011 |title=Habitat preferences and activity patterns of the larger mammal community in Phnom Prich Wildlife Sanctuary, Cambodia |journal=The Raffles Bulletin of Zoology |volume=59 |issue=2 |pages=311−318 |url=https://www.researchgate.net/publication/228843849 |access-date=2019-04-16 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150044/https://www.researchgate.net/publication/228843849_Habitat_preferences_and_activity_patterns_of_the_larger_mammal_community_in_Phnom_Prich_Wildlife_Sanctuary_Cambodia |url-status=live}}</ref><ref>{{cite journal |last1=Gray |first1=T. N. E. |year=2013 |title=Activity patterns and home ranges of Indochinese leopard ''Panthera pardus delacouri'' in the Eastern Plains Landscape, Cambodia |journal=Natural History Bulletin of the Siam Society |volume=59 |pages=39−47 |url=http://www.siamese-heritage.org/nhbsspdf/vol051-060/NHBSS_059_1h_Gray_ActivityPatternsAndH.pdf |archive-url=https://web.archive.org/web/20160222060912/http://www.siamese-heritage.org/nhbsspdf/vol051-060/NHBSS_059_1h_Gray_ActivityPatternsAndH.pdf |archive-date=2016-02-22 |url-status=live}}</ref>
In southern China, leopards were recorded only in the [[Qinling]] Mountains during surveys in 11 nature reserves between 2002 and 2009.<ref>{{cite journal |author=Li, S. |author2=Wang, D. |author3=Lu, Z. |author4=Mc Shea, W. J. |name-list-style=amp |year=2010 |title=Cats living with pandas: The status of wild felids within giant panda range, China |journal=Cat News |volume=52 |pages=20–23}}</ref>
In Java, leopards inhabit dense [[tropical rainforest]]s and dry [[deciduous forest]]s at elevations from sea level to {{cvt|2540|m}}. Outside protected areas, leopards were recorded in mixed [[agricultural]] land, [[secondary forest]] and [[Agroforestry|production forest]] between 2008 and 2014.<ref>{{cite journal |last1=Wibisono |first1=H. T. |last2=Wahyudi |first2=H. A. |last3=Wilianto |first3=E. |last4=Pinondang |first4=I. M. R. |last5=Primajati |first5=M. |last6=Liswanto |first6=D. |last7=Linkie |first7=M. |name-list-style=amp |year=2018 |title=Identifying priority conservation landscapes and actions for the Critically Endangered Javan leopard in Indonesia: Conserving the last large carnivore in Java Island |journal=PLOS ONE |volume=13 |issue=6 |page=e0198369 |doi=10.1371/journal.pone.0198369 |pmid=29949588 |pmc=6021038 |bibcode=2018PLoSO..1398369W |doi-access=free}}</ref>
In the Russian Far East, it inhabits [[temperate coniferous forest]]s where winter temperatures reach a low of {{cvt|−25|°C|°F}}.<ref name=Uphyrkina/>
== Behaviour and ecology ==
The leopard is a solitary and [[Territory (animal)|territorial]] animal. It is typically shy and alert when crossing roadways and encountering oncoming vehicles, but may be emboldened to attack people or other animals when threatened. Adults associate only in the mating season. Females continue to interact with their offspring even after weaning and have been observed sharing kills with their offspring when they can not obtain any prey. They produce a number of vocalizations, including growls and snarls. Cubs call their mother with meows and an ''urr-urr'' sound.<ref name=estes/> The most notable vocalization is the 'sawing' [[roar]], which consists of deep, repeated strokes. This likely functions in establishing territories and attracting mates.<ref>{{cite journal|last1=Growcott|first1=J|last2=Lobora|first2=A|last3=Markham|first3=A|last4=Searle|first4=C. E.|last5=Wahlström|first5=J|last6=Wijers|first6=M|last7=Simmons|first7=B. I.|year=2024| title=The secret acoustic world of leopards: A paired camera trap and bioacoustics survey facilitates the individual identification of leopards via their roars|journal=Remote Sensing in Ecology and Conservation|article-number=rse2.429|doi=10.1002/rse2.429}}</ref>
The whitish spots on the back of its ears are thought to play a role in communication.<ref>{{cite book |last1=Leyhausen |first1=P. |year=1979 |title=Cat behavior: the predatory and social behavior of domestic and wild cats |___location=Berlin |publisher=Garland Publishing, Incorporated |page=281 |isbn=9780824070175}}</ref>
It has been hypothesized that the white tips of their tails may function as a 'follow-me' signal in [[intraspecific]] communication. However, no significant association were found between a conspicuous colour of tail patches and behavioural variables in carnivores.<ref>{{cite journal |last1=Ortolani |first1=A. |year=1999 |title=Spots, stripes, tail tips and dark eyes: predicting the function of carnivore colour patterns using the comparative method |journal=Biological Journal of the Linnean Society |volume=67 |issue=4 |pages=433–476 |doi=10.1111/j.1095-8312.1999.tb01942.x |doi-access=free}}</ref><ref>{{cite journal |last1=Caro |first1=T. |year=2005 |title=The adaptive significance of coloration in mammals |journal=BioScience |volume=55 |issue=2 |pages=125–136 |doi=10.1641/0006-3568(2005)055[0125:TASOCI]2.0.CO;2 |doi-access=free}}</ref>
Leopards are mainly active from dusk till dawn and will rest for most of the day and some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking {{cvt|1|-|25|km|mi}} across their range at night; wandering up to {{cvt|75|km}} if disturbed.<ref name=estes/><ref name=nowak/> In some regions, they are [[Nocturnality|nocturnal]].<ref name=Hunter_al2003>{{cite journal |last1=Hunter |first1=L. |last2=Balme |first2=G. |last3=Walker |first3=C. |last4=Pretorius |first4=K. |last5=Rosenberg |first5=K. |name-list-style=amp |year=2003 |title=The landscape ecology of leopards (''Panthera pardus'') in northern KwaZulu-Natal, South Africa: a preliminary project report |journal=Ecological Journal |volume=5 |pages=24–30 |url=http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf |url-status=dead |archive-url=https://web.archive.org/web/20090304233241/http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf |archive-date=March 4, 2009}} {{open access}}</ref><ref>{{cite journal |last1=Spalton |first1=J.A. |name-list-style=amp |last2=Al Hikmani |first2=H. M. |last3=Willis |first3=D. |last4=Said |first4=A. S. B. |title=Critically endangered Arabian leopards ''Panthera pardus nimr'' persist in the Jabal Samhan Nature Reserve, Oman |journal=Oryx |date=2006 |volume=40 |issue=3 |pages=287–294 |doi=10.1017/S0030605306000743 |doi-access=free}}</ref> In western African forests, they have been observed to be largely [[Diurnality|diurnal]] and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.<ref>{{cite journal |last1=Jenny |first1=D. |last2=Zuberbuhler |first2=K. |name-list-style=amp |title=Hunting behaviour in west African forest leopards |journal=African Journal of Ecology |date=2005 |volume=43 |issue=3 |pages=197–200 |doi=10.1111/j.1365-2028.2005.00565.x |bibcode=2005AfJEc..43..197J |url=https://www.researchgate.net/publication/229882994 |access-date=2018-09-25 |archive-date=2018-09-25 |archive-url=https://web.archive.org/web/20180925142137/https://www.researchgate.net/publication/229882994 |url-status=live}}</ref>
Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.<ref name=CAP/>
They can run at over {{cvt|58|km/h|mph m/s}}, leap over {{cvt|6|m}} horizontally, and jump up to {{cvt|3|m}} vertically.<ref name="WCW">{{Cite book |last1=Sunquist |first1=M. E. |year=2002 |last2=Sunquist |first2=F. |name-list-style=amp |title=Wild Cats of the World |publisher=University of Chicago Press |___location=Chicago |isbn=978-0-226-77999-7 |chapter=Leopard ''Panthera pardus'' |pages=318–342 |chapter-url=https://books.google.com/books?id=IF8nDwAAQBAJ&pg=PA320 |access-date=2021-05-19 |archive-date=2024-04-30 |archive-url=https://web.archive.org/web/20240430033051/https://books.google.com/books?id=IF8nDwAAQBAJ&pg=PA320#v=onepage&q&f=false |url-status=live}}</ref>
{{image frame|align=center|border=no|content=
<gallery mode=packed>
leopard rear view soft.jpg|A female leopard with white markings on the backs of her ears
leopard walking.jpg|A female leopard showing white spots on the tail
Leopard davidraju 68.jpg|A leopard climbing down a tree
Leopard hunting a bush pig - DPLA - 57da78c992bc6073d2751f3f8936aad0.jpg|A leopard hunting a [[bushpig]]
Leopard.ogv|Video of a leopard in the wild
</gallery>
}}
===
[[File:Day 47 Leopard (Panthera pardus) male marking a bush with urine ... (53310719953).jpg|thumb|A male leopard [[scent mark|scent-marking]] his territory<ref>{{Cite book |last1=Bothma |first1=Jacobus du P. |url=https://books.google.com/books?id=DPvuCAAAQBAJ |title=Larger Carnivores of the African Savannas |last2=Walker |first2=Clive |date=2013-11-11 |publisher=Springer Science & Business Media |isbn=978-3-662-03766-9 |language=en |access-date=2024-01-06 |archive-date=2024-01-06 |archive-url=https://web.archive.org/web/20240106170223/https://books.google.com/books?id=DPvuCAAAQBAJ |url-status=live}}</ref>]]
In [[Kruger National Park]], most leopards tend to keep {{cvt|1|km|mi}} apart.<ref name=bailey93>{{cite book |last=Bailey |first=T. N. |year=1993 |title=The African leopard: a study of the ecology and behaviour of a solitary felid |publisher=Columbia University Press |___location=New York |isbn=978-1-932846-11-9}}</ref> Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.<ref name="Kingdon"/><ref>{{Cite journal |last1=Pirie |first1=T. J.|last2=Thomas |first2=R. L.|last3=Reilly|first3=B. K. |last4=Fellowes |first4=M. D. E. |name-list-style=amp |date=2014 |title=Social interactions between a male leopard (''Panthera pardus'') and two generations of his offspring |journal=African Journal of Ecology |volume=52 |issue=4 |pages=574–576 |doi=10.1111/aje.12154 |doi-access=free |bibcode=2014AfJEc..52..574P}}</ref> Aggressive encounters are rare, typically limited to defending territories from intruders.<ref name=mammal/> In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.<ref name=Hunter_al2003/>
Males occupy [[home range]]s that often overlap with a few smaller female home ranges, probably as a strategy to enhance access to females. In the [[Ivory Coast]], the home range of a female was completely enclosed within a male's.<ref name=Ivory>{{cite journal |author=Jenny, D. |year=1996 |title=Spatial organization of leopards ''Panthera pardus'' in Tai National Park, Ivory Coast: Is rainforest habitat a "tropical haven"? |journal=Journal of Zoology |issue=3 |volume=240 |pages=427–440 |doi=10.1111/j.1469-7998.1996.tb05296.x |url=https://www.researchgate.net/publication/227901726 |access-date=2021-09-06 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505145917/https://www.researchgate.net/publication/227901726_Spatial_organization_of_leopards_Panthera_pardus_in_Tai_National_Park_Ivory_Coast_Is_rainforest_habitat_a_%27tropical_haven%27 |url-status=live}}</ref> Females live with their cubs in home ranges that overlap extensively, probably due to the association between mothers and their offspring. There may be a few other fluctuating home ranges belonging to young individuals. It is not clear if male home ranges overlap as much as those of females do. Individuals try to drive away intruders of the same sex.<ref name=estes/><ref name=nowak/>
A study of leopards in the Namibian farmlands showed that the size of home ranges was not significantly affected by sex, rainfall patterns or season; the higher the prey availability in an area, the greater the leopard population density and the smaller the size of home ranges, but they tend to expand if there is human interference.<ref>{{cite journal|last1=Marker |first1=L. L. |last2=Dickman |first2=A. J. |name-list-style=amp |title=Factors affecting leopard (''Panthera pardus'') spatial ecology, with particular reference to Namibian farmlands |journal=South African Journal of Wildlife Research |date=2005 |volume=35 |issue=2 |pages=105–115 |url=http://reference.sabinet.co.za/webx/access/journal_archive/03794369/3418.pdf}} {{open access}}</ref>
Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the [[Serengeti]], males have home ranges of {{cvt|33|-|38|km2|sqmi}} and females of {{cvt|14|-|16|km2|sqmi}};<ref>{{cite journal |last1=Bertram |first1=B. C. R.|title=Leopard ecology as studied by radio tracking |journal=Symposia of the Zoological Society of London |date=1982 |volume=49|pages=341–352}}</ref><ref>{{cite journal |last1=Mizutani, F. |last2=Jewell, P. A. |name-list-style=amp |year=1998 |title=Home-range and movements of leopards (''Panthera pardus'') on a livestock ranch in Kenya |journal=Journal of Zoology |pages=269–286 |volume=244 |issue=2 |doi=10.1017/S0952836998002118}}</ref> but males in northeastern Namibia of {{cvt|451|km2|sqmi}} and females of {{cvt|188|km2|sqmi}}.<ref>{{cite journal |last1=Stander |first1=P. E. |last2=Haden |first2=P. J. |last3=Kaqece |first3=II. |last4=Ghau |first4=II. |name-list-style=amp|title=The ecology of asociality in Namibian leopards|journal=Journal of Zoology |date=1997 |volume=242|issue=2 |pages=343–364 |doi=10.1111/j.1469-7998.1997.tb05806.x}}</ref> They are even larger in arid and montane areas.<ref name=mammal/> In Nepal's [[Bardia National Park]], male home ranges of {{cvt|48|km2|sqmi}} and female ones of {{cvt|5|-|7|km2|sqmi}} are smaller than those generally observed in Africa.<ref>{{cite journal |author=Odden, M. |author2=Wegge, P. |name-list-style=amp |year=2005 |title=Spacing and activity patterns of leopards ''Panthera pardus'' in the Royal Bardia National Park, Nepal |journal=Wildlife Biology |volume=11 |issue=2 |pages=145–152 |doi=10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2 |doi-access=free |s2cid=86140708}}</ref>
==
{{multiple image |perrow=1 |image1=Leopard (Panthera pardus pardus) stalking.jpg |caption1=Leopard stalking |image2=Leopard kill - KNP - 001.jpg|caption2=Leopard applying a throat bite to a [[Cape bushbuck|bushbuck]]|image3=Leopardo (Panthera pardus) devorando un antílope, parque nacional Kruger, Sudáfrica, 2018-07-26, DD 06.jpg |caption3=Leopard caches a kill in a tree}}
The leopard is a [[carnivore]] that prefers medium-sized prey with a body mass ranging from {{cvt|10|–|40|kg|lb}}. Prey species in this weight range tend to occur in dense habitat and to form small herds. Species that prefer open areas and have well-developed anti-predator strategies are less preferred. More than 100 prey species have been recorded. The most preferred species are [[ungulates]], such as [[impala]], [[Cape bushbuck|bushbuck]], [[common duiker]] and [[chital]]. [[Primate]]s preyed upon include [[white-eyelid mangabey]]s, [[guenon]]s and [[gray langur]]s. Leopards also kill smaller carnivores like [[black-backed jackal]], [[bat-eared fox]], [[genet (animal)|genet]] and cheetah.<ref name=Hayward2006>{{cite journal |last1=Hayward |first1=M.W. |last2=Henschel |first2=P. |last3=O'Brien |first3=J. |last4=Hofmeyr |first4=M. |last5=Balme |first5=G. |last6=Kerley |first6=G. I. H. |name-list-style=amp |title=Prey preferences of the leopard (''Panthera pardus'') |journal=Journal of Zoology |date=2006 |volume=270 |issue=4 |pages=298–313 |doi=10.1111/j.1469-7998.2006.00139.x |url=http://www.ibs.bialowieza.pl/g2/pdf/1596.pdf |archive-url=https://web.archive.org/web/20121105063845/http://www.ibs.bialowieza.pl/g2/pdf/1596.pdf |archive-date=2012-11-05 |url-status=live}}</ref> In urban environments, [[domestic dog]]s provide an important food source.<ref name=Powell2021/> The largest prey killed by a leopard was reportedly a male [[Taurotragus|eland]] weighing {{cvt|900|kg}}.<ref name=WCW/> A study in [[Wolong National Nature Reserve]] in southern China demonstrated variation in the leopard's diet over time; over the course of seven years, the vegetative cover receded, and leopards opportunistically shifted from primarily consuming [[tufted deer]] to pursuing [[Chinese bamboo rat|bamboo rat]]s and other smaller prey.<ref>{{cite journal |author1=Johnson, K. G. |author2=Wei, W. |author3=Reid, D. G. |author4=Jinchu, H. |date=1993 |title=Food habits of Asiatic leopards (''Panthera pardus fusca'') in Wolong Reserve, Sichuan, China |journal=Journal of Mammalogy |volume=74 |issue=3 |pages=646–650 |jstor=1382285 |doi=10.2307/1382285}}</ref>
The leopard depends mainly on its acute senses of hearing and vision for hunting.<ref name=Mills>{{cite book |last1=Mills |first1=M. G. L. |last2=Hes |first2=L. |name-list-style=amp |isbn=978-0-947430-55-9|pages=178–180 |publisher=Struik Publishers|___location=Cape Town, South Africa |title=The Complete Book of Southern African Mammals |year=1997}}</ref> It primarily hunts at night in most areas.<ref name=estes/> In western African forests and Tsavo National Park, they have also been observed hunting by day.<ref name=hamilton76>{{cite thesis |last=Hamilton |first=P. H. |year=1976 |title=The movements of leopards in Tsavo National Park, Kenya, as determined by radio-tracking |degree= M.Sc. |publisher=University of Nairobi |___location=Nairobi}}</ref> They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.<ref name= Kruuk>{{cite journal |last1=Kruuk |first1=H. |last2=Turner |first2=M. |name-list-style=amp |date=1967 |title=Comparative notes on predation by lion, leopard, cheetah and wild dog in the Serengeti area, East Africa |journal=Mammalia |volume=31 |issue=1 |pages=1–27 |doi=10.1515/mamm.1967.31.1.1 |s2cid=84619500}}</ref> It stalks its prey and tries to approach as closely as possible, typically within {{cvt|5|m}} of the target, and, finally, pounces on it and kills it by suffocation. It kills small prey with a bite to the back of the neck, but holds larger animals [[Throat clamp|by the throat and strangles them]].<ref name=estes/> It [[Cache (biology)|caches]] kills up to {{cvt|2|km|mi}} apart.<ref name=Kingdon/> It is able to take large prey due to its powerful jaw muscles, and is therefore strong enough to drag carcasses heavier than itself up into trees; an individual was seen to haul a young giraffe weighing nearly {{cvt|125|kg}} up {{cvt|5.7|m|ftin}} into a tree.<ref name=hamilton76/> It eats small prey immediately, but drags larger carcasses over several hundred metres and caches it safely in trees, bushes or even caves; this behaviour allows the leopard to store its prey away from rivals, and offers it an advantage over them. The way it stores the kill depends on local topography and individual preferences, varying from trees in Kruger National Park to bushes in the plain terrain of the Kalahari.<ref name=mammal/><ref name=Schaller72>{{cite book |last=Schaller |first=G. |year=1972 |title=Serengeti: a kingdom of predators |publisher=Knopf |___location=New York |isbn=978-0-394-47242-3 |url=https://books.google.com/books?id=wJ_wAAAAMAAJ |access-date=2020-09-18 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150524/https://books.google.com/books?id=wJ_wAAAAMAAJ |url-status=live}}</ref> Before their extirpation from Europe, leopards there cached their meat in caves, as evidenced by fossilised bone accumulations in caves such as Los Rincones in the [[Province of Zaragoza]], Spain.<ref>{{Cite journal |last1=Sauqué |first1=Víctor |last2=Rabal-Garcés |first2=Raquel |last3=Sola-Almagro |first3=Cristina |last4=Cuenca-Bescós |first4=Gloria |date=18 March 2014 |editor-last=Rook |editor-first=Lorenzo |title=Bone Accumulation by Leopards in the Late Pleistocene in the Moncayo Massif (Zaragoza, NE Spain) |journal=[[PLoS ONE]] |language=en |volume=9 |issue=3 |pages=e92144 |doi=10.1371/journal.pone.0092144 |pmid=24642667 |pmc=3958443 |bibcode=2014PLoSO...992144S |doi-access=free |issn=1932-6203 }}</ref>
Average daily consumption rates of {{cvt|3.5|kg|lboz}} were estimated for males and of {{cvt|2.8|kg|lboz}} for females.<ref name=bailey93/> In the southern [[Kalahari Desert]], leopards meet their water requirements by the bodily fluids of prey and [[succulent plant]]s; they drink water every two to three days and feed infrequently on moisture-rich plants such as [[gemsbok cucumber]]s, [[watermelon]] and Kalahari [[Schmidtia|sour grass]].<ref>{{cite journal| last1=Bothma |first1=J. du P. |title=Water-use by southern Kalahari leopards |journal=South African Journal of Wildlife Research |date=2005 |volume=35 |pages=131–137 |url=http://reference.sabinet.co.za/webx/access/journal_archive/03794369/2353.pdf}} {{open access}}</ref>
===
[[File:Lioness vs Leopard 9 July 2016 Latest Sightings 1.png|thumb|A lioness steals a leopard kill in Kruger National Park]]
Across its range, the leopard coexists with a number of other large predators. In Africa, it is part of a large predator [[Guild (ecology)|guild]] with lions, cheetahs, [[spotted hyena|spotted]] and [[brown hyena]]s, and [[African wild dog]]s.<ref name=Rafiaq2020>{{cite journal|last1=Rafiq|first1=K.|last2=Wayward |first2=M. W.|last3=Wilson|first3=A. M.|last4=Meloro|first4=C. |last5=Jordan|first5=N. R. |last6=Wich |first6=S. A. |last7=McNutt|first7=J. W. |last8=Golabek|first8=K. A.|year=2020|title=Spatial and temporal overlaps between leopards (''Panthera pardus'') and their competitors in the African large predator guild|journal=Journal of Zoology |volume=311 |issue=4 |pages=246–259 |doi=10.1111/jzo.12781}}</ref> The leopard is dominant only over the cheetah while the others have the advantage of size, pack numbers or both.<ref name=estes/> Lions pose a great mortal threat and can be responsible for 22% of leopard deaths in [[Sabi Sand Game Reserve]]. Spotted hyenas are less threatening but are more likely to [[Kleptoparasitism|steal kills]], being the culprits of up to 50% of stolen leopard kills in the same area.<ref>{{cite journal|last1=Balme|first1=G. A.|last2=Pitman|first2=R. T.|last3=Robinson|first3=H. S.|last4=Miller|first4=J. R. B.|last5=Funston|first5=P. J.|last6=Hunter|first6=L. T. B.|year=2017|title=Leopard distribution and abundance is unaffected by interference competition with lions|journal=Behavioral Ecology |volume=28 |issue=5 |pages=1348–1358 |doi=10.1093/beheco/arx098}}</ref><ref name=Balme2017>{{cite journal|last1=Balme|first1=G. A. |last2=Miller |first2=J. R. B.|last3=Pitman|first3=R. T.|last4=Hunter|first4=L. T. B.|year=2017|title=Caching reduces kleptoparasitism in a solitary, large felid|journal=Journal of Animal Ecology|volume=86|issue=3 |pages=634–644 |doi=10.1111/1365-2656.12654 |pmid=28217865 |bibcode=2017JAnEc..86..634B}}</ref> To counter this, leopards store their kills in the trees and out of reach.<ref name=Balme2017/><ref name=Vissia2022>{{cite journal|last1=Vissia|first1=S.|last2=Fattebert|first2=J. |last3=van Langevelde|first3=F. |year=2022 |title=Leopard density and interspecific spatiotemporal interactions in a hyena-dominated landscape|journal=Ecology and Evolution |volume=12 |issue=10 |page=e9365 |doi=10.1002/ece3.9365 |pmid=36225822 |pmc=9534747 |bibcode=2022EcoEv..12E9365V}}</ref> Lions have a high success rate in fetching leopard kills from trees.<ref name=Balme2017/> Leopards do not seem to actively avoid their competitors but rather difference in prey and habitat preferences appear to limit their spatial overlap.<ref name=Rafiaq2020/> In particular, leopards use heavy vegetation regardless of whether lions are present in an area and both cats are active at the same time of day.<ref>{{cite journal |last1=Miller|first1=J. R. B. |last2=Pitman |first2=R. T. |last3=Mann |first3=G. K. H. |last4=Fuller|first4=A. K.|last5=Balme|first5=G. A. |year=2018|title=Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition |journal=Journal of Animal Ecology |volume=87 |issue=6 |pages=1709–1726 |doi=10.1111/1365-2656.12883 |pmid=30010193 |bibcode=2018JAnEc..87.1709M}}</ref>
In Asia, the leopard's main competitors are tigers and [[dhole]]s. Both the larger tiger and pack-living dhole dominate leopards during encounters. Interactions between the three predators involve chasing, stealing kills and direct killing.<ref>{{cite journal |last1=Srivathsa |first1=A. |last2=Ramachandran|first2=V. |last3=Saravanan |first3=P. |last4=Sureshbabu |first4=A. |last5=Ganguly |first5=D. |last6=Ramakrishnan |first6=U. |year=2023 |title=Topcats and underdogs: intraguild interactions among three apex carnivores across Asia's forestscapes |journal=Biological Reviews |volume=98 |issue=6 |pages=2114–2135 |doi=10.1111/brv.12998 |pmid=37449566 |s2cid=259903849 |name-list-style=amp}}</ref> Tigers appear to inhabit the deep parts of the forest while leopards and dholes are pushed closer to the fringes.<ref>{{cite journal |author=Thinley, P. |author2=Rajaratnam, R. |author3=Lassoie, J. P. |author4=Morreale, S. J. |author5=Curtis, P. D. |author6=Vernes, K. |author7=Leki Leki |author8=Phuntsho, S. |author9=Dorji, T. |author10=Dorji, P. |name-list-style=amp |year=2018 |title=The ecological benefit of tigers (''Panthera tigris'') to farmers in reducing crop and livestock losses in the eastern Himalayas: Implications for conservation of large apex predators |journal=Biological Conservation |volume=219 |pages=119–125 |doi=10.1016/j.biocon.2018.08.007 |doi-access=free}}</ref> The three predators coexist by hunting different sized prey.<ref name=Karanth>{{cite journal |last=Karanth |first=K. U. |author2=Sunquist, M. E. |name-list-style=amp |year=2000 |title=Behavioural correlates of predation by tiger (''Panthera tigris''), leopard (''Panthera pardus'') and dhole (''Cuon alpinus'') in Nagarahole, India | journal=Journal of Zoology |volume=250 |pages=255–265 |doi=10.1111/j.1469-7998.2000.tb01076.x |issue=2}}</ref> In [[Nagarhole National Park]], the average size for a leopard kill was {{cvt|37.6|kg}} compared to {{cvt|91.5|kg}} for tigers and {{cvt|43.4|kg}} for dholes.<ref name=KaranthSunquist1995>{{cite journal |author1=Karanth, K. U. |author2=Sunquist, M. E. |name-list-style=amp |title=Prey selection by tiger, leopard and dhole in tropical forests |jstor=5647 |doi=10.2307/5647 |journal=Journal of Animal Ecology |volume=64 |issue=4 |year=1995 |pages=439–450|bibcode=1995JAnEc..64..439K}}</ref> At [[Kui Buri National Park]], following a reduction in prey numbers, tigers continued to feed on favoured prey while leopards and dholes had to increase their consumption of small prey.<ref>{{cite journal |author1=Steinmetz, R. |name-list-style=amp |author2=Seuaturien, N. |author3=Intanajitjuy, P. |author4=Inrueang, P. |author5=Prempree, K. |year=2021 |title=The effects of prey depletion on dietary niches of sympatric apex predators in Southeast Asia |journal=Integrative Zoology |volume=16 |issue=1 |pages=19–32 |doi=10.1111/1749-4877.12461|pmid=32627329 }}</ref> Leopards can live successfully in tiger habitat when there is abundant food and vegetation cover.<ref name=Karanth/><ref>{{cite journal |author1=Odden, M. |name-list-style=amp |author2=Wegge, P. |author3=Fredriksen, T. |year=2010 |title=Do tigers displace leopards? If so, why? |journal=Ecological Research |volume=25 |issue=4 |pages=875–881 |doi=10.1007/s11284-010-0723-1 |bibcode=2010EcoR...25..875O}}</ref> Otherwise, they appear to be less common where tigers are numerous. The recovery of the tiger population in [[Rajaji National Park]] during the 2000s led to a reduction in leopard population densities.<ref>{{cite journal |last1=Harihar |first1=A. |last2=Pandav |first2=B. |last3=Goyal |first3=S. P. |year=2011 |title=Responses of leopard ''Panthera pardus'' to the recovery of a tiger ''Panthera tigris'' population |journal=Journal of Applied Ecology |volume=48 |issue=3 |pages=806–814 |doi=10.1111/j.1365-2664.2011.01981.x|doi-access=free |bibcode=2011JApEc..48..806H |name-list-style=amp}}</ref>
===Reproduction and life cycle===
{{multiple image |perrow=1 |image2=Leopards mating.jpg |caption2=Leopards mating |image3=David Raju Leopard 3457 (cropped).jpg |caption3=Leopard cubs in tree}}
In some areas, leopards mate all year round. In [[Manchuria]] and [[Siberia]], they mate during January and February. On average, females begin to breed between the ages of 2½ and three, and males between the ages of two and three.<ref name=CAP/> The female's [[estrous cycle]] lasts about 46 days, and she is usually in heat for 6–7 days.<ref>{{cite journal |doi=10.1111/j.1748-1090.1966.tb01746.x |author=Sadleir, R. |year=1966 |title=Notes on the Reproduction of the larger Felidae |journal=International Zoo Yearbook|volume=6 |pages=184–187}}</ref> [[Gestation]] lasts for 90 to 105 days.<ref>{{cite book |author=Hemmer, H. |year=1976 |chapter=Gestation period and postnatal development in felids |pages=143–165 |editor=Eaton, R.L. |title=The World's Cats |volume=3 |publisher=Carnivore Research Institute, Univ. Washington, Seattle}}</ref> Cubs are usually born in a [[Litter (zoology)|litter]] of 2–4 cubs.<ref>{{cite journal |author=Eaton, R.L. |year=1977 |title=Reproductive biology of the leopard |journal=Zoologischer Garten |volume=47 |issue=5 |pages =329–351}}</ref> The mortality rate of cubs is estimated at 41–50% during the first year.<ref name="bailey93" /> Predators are the biggest cause for leopard cub mortality during their first year. Male leopards are known to cause [[infanticide (zoology)|infanticide]], in order to bring the female back into heat.<ref name=Kingdon/> Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.<ref name=CAP/>
Females give birth in a cave, crevice among boulders, hollow tree or thicket. Newborn cubs weigh {{Cvt|280-1000|g}}, and are born with closed eyes, which open four to nine days after birth.<ref name=WCW/><ref name=mammal/> The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in colour with less defined spots. They begin to eat meat at around nine weeks.<ref name="Kingdon"/> Around three months of age, the young begin to follow the mother on hunts. At one year of age, cubs can probably fend for themselves, but will remain with the mother for 18–24 months.<ref name=CMBC>{{cite web |url=https://brainmuseum.org/Specimens/carnivora/leopard/index.html |title=Leopard (''Panthera pardus''); Physical characteristics and distribution |work=Comparative Mammalian Brain Collections |access-date=2008-06-07 |archive-date=2011-01-03 |archive-url=https://web.archive.org/web/20110103183140/http://www.brainmuseum.org/specimens/carnivora/leopard/index.html |url-status=live}}</ref> After separating from their mother, sibling cubs may travel together for months.<ref name=CAP/> Both male and female leopards typically reach sexual maturity at 2–2⅓ years.<ref name=Kingdon/>
The [[Generation time|generation length]] of the leopard is 9.3 years.<ref>{{cite journal |title=Generation length for mammals |last1=Pacifici |first1=M. |last2=Santini |first2=L. |last3=Di Marco |first3=M. |last4=Baisero |first4=D. |name-list-style=amp |last5=Francucci |first5=L. |last6=Grottolo Marasini |first6=G. |last7=Visconti |first7=P. |last8=Rondinini |first8=C. |journal=Nature Conservation |year=2013 |issue=5 |pages=87–94}}</ref>
The average life span of a leopard is 12–17 years.<ref name=WCW/>
The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.<ref>{{cite news |author=Salisbury, S. |date=2014 |url=http://www.palmbeachpost.com/news/news/roxanne-oldest-spotted-leopard-in-captivity-dies-a/ngyYc/ |title=Roxanne, oldest spotted leopard in captivity, dies at Acreage preserve |newspaper=The Palm Beach Post |archive-date=2014-08-11|archive-url=https://web.archive.org/web/20140811165340/http://www.palmbeachpost.com/news/news/roxanne-oldest-spotted-leopard-in-captivity-dies-a/ngyYc/}}</ref>
== Conservation ==
The leopard is listed on [[CITES Appendix I]], and hunting is banned in Botswana and Afghanistan; in 11 sub-Saharan countries, trade is restricted to skins and body parts of 2,560 individuals.<ref name=IUCN/>
In 2007, a leopard reintroduction programme was initiated in the Russian Caucasus, where captive bred individuals are reared and trained in {{cvt|0.5–0.9|ha}} large enclosures in [[Sochi National Park]]; six individuals released into [[Caucasus Nature Reserve]] and [[Alaniya National Park]] in 2018 survived as of February 2022.<ref>{{cite journal |author1=Rozhnov, V.V. |name-list-style=amp |author2=Yachmennikova, A.A. |author3=Dronova, N.A. |author4=Naidenko, S.V. |author5=Hernandez-Blanco, J.A. |author6=Chistopolova, M.D. |author7=Pkhitikov, A.B. |author8=Tembotova, F.A. |author9=Trepet, S.A. |author10=Chestin, I.E. |year=2022 |title=Experience of the leopard re-covering through reintroduction in the Russian Caucasus |journal=Cat News |issue=Special issue |pages=67–71 |url=http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.14._SI_15/Cat_News_Special_Issue_15_Persian_leopard_FINAL.pdf#page=67 |access-date=2023-11-11 |archive-date=2023-11-11 |archive-url=https://web.archive.org/web/20231111083835/http://www.catsg.org/fileadmin/filesharing/5.Cat_News/5.3._Special_Issues/5.3.14._SI_15/Cat_News_Special_Issue_15_Persian_leopard_FINAL.pdf#page=67 |url-status=live}}</ref>
=== Threats===
The leopard is primarily threatened by [[habitat fragmentation]] and conversion of forest to agriculturally used land, which lead to a declining natural prey base, [[human–wildlife conflict]] with livestock herders and high leopard mortality rates. It is also threatened by [[trophy hunting]] and [[poaching]].<ref name=IUCN /> Contemporary records suggest that the leopard occurs in only 25% of its historical range.<ref name=Jacobson2016/><ref name=Williams2017>{{cite journal|last1=Williams|first1=S. T. |last2=Williams |first2=K. S. |last3=Lewis |first3=B. P. |last4=Hill |first4=R. A. |name-list-style=amp |title=Population dynamics and threats to an apex predator outside protected areas: implications for carnivore management |journal=Royal Society Open Science |date=2017 |volume=4 |issue=4 |pages=161090 |doi=10.1098/rsos.161090 |doi-access=free |pmid=28484625 |pmc=5414262 |bibcode=2017RSOS....461090W}}</ref>
Between 2002 and 2012, at least four leopards were estimated to have been poached per week in India for the [[illegal wildlife trade]] of its skins and bones.<ref>{{Cite report |author1=Raza, R.H. |name-list-style=amp |author2=Chauhan, D.S. |author3=Pasha, M.K.S. |author4=Sinha, S. |date=2012 |title=Illuminating the blind spot: A study on illegal trade in Leopard parts in India (2001–2010) |publisher=TRAFFIC India, WWF India |___location=New Delhi |url=https://www.traffic.org/site/assets/files/4130/illuminating_the_blind_spot.pdf |archive-url=https://web.archive.org/web/20200924143713/https://www.traffic.org/site/assets/files/4130/illuminating_the_blind_spot.pdf |archive-date=2020-09-24 |url-status=live}}</ref>
In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.<ref name=Bergin>{{cite journal |last1=Bergin |first1=D. |last2=Nijman |first2=V. |name-list-style=amp |year=2014 |title=Open, Unregulated Trade in Wildlife in Morocco's Markets |journal=TRAFFIC Bulletin |volume=26 |issue=1 |pages=65–70 |url=https://www.researchgate.net/publication/267748463 |access-date=2015-03-23 |archive-date=2018-10-31 |archive-url=https://web.archive.org/web/20181031133219/https://www.researchgate.net/publication/267748463 |url-status=live}}</ref> In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.<ref>{{Cite journal |last1=Bergin |first1=D. |last2=Nijman |first2=V. |name-list-style=amp |date=2015 |title=Potential benefits of impending Moroccan wildlife trade laws, a case study in carnivore skins |journal=Biodiversity and Conservation |volume=25 |issue=1 |pages=199–201 |doi=10.1007/s10531-015-1042-1 |s2cid=34533018 |url=https://www.researchgate.net/publication/287807462 |access-date=2017-01-11 |archive-date=2020-01-07 |archive-url=https://web.archive.org/web/20200107220433/https://www.researchgate.net/publication/287807462_Potential_benefits_of_impending_Moroccan_wildlife_trade_laws_a_case_study_in_carnivore_skins |url-status=live}}</ref>
Surveys in the Central African Republic's [[Chinko]] area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period, [[transhumant]] [[Pastoralism|pastoralist]]s from the border area with Sudan moved in the area with their livestock. Rangers confiscated large amounts of [[poison]] in the camps of livestock herders who were accompanied by armed merchants. They engaged in poaching large herbivores, sale of [[bushmeat]] and trading leopard skins in [[Am Dafok]].<ref>{{cite journal |author1=Äbischer, T. |author2=Ibrahim, T. |author3=Hickisch, R. |author4=Furrer, R. D. |name-list-style=amp |author5=Leuenberger, C. |author6=Wegmann, D. |year=2020 |title=Apex predators decline after an influx of pastoralists in former Central African Republic hunting zones |journal=Biological Conservation |volume=241 |article-number=108326 |doi=10.1016/j.biocon.2019.108326 |bibcode=2020BCons.24108326A |s2cid=213766740 |url=https://doc.rero.ch/record/328409/files/weg_apd.pdf |archive-url=https://web.archive.org/web/20201003124917/https://doc.rero.ch/record/328409/files/weg_apd.pdf |archive-date=2020-10-03 |url-status=live}}</ref>
In Java, the leopard is threatened by illegal hunting and trade. Between 2011 and 2019, body parts of 51 Javan leopards were seized including six live individuals, 12 skins, 13 skulls, 20 canines and 22 claws.<ref>{{cite journal |author1=Gomez, L. |name-list-style=amp |author2=Shepherd, C.R. |year=2021 |title=The illegal exploitation of the Javan Leopard (''Panthera pardus melas'') and Sunda Clouded Leopard (''Neofelis diardi'') in Indonesia |journal=Nature Conservation |volume=43 |issue=43 |pages=25–39 |doi=10.3897/natureconservation.43.59399 |doi-access=free |s2cid=233286106}}</ref>
==
===Cultural significance===
{{Multiple image|direction=vertical|align=right|image1=Sousse museum Smirat-retouched.jpg|caption1=Leopards on the [[Magerius Mosaic]] from modern [[Tunisia]] |image2=Leopard head hip ornament, Nigeria, Court of Benin, Edo people, late 18th century, bronze, copper, iron - De Young Museum - DSC01037.JPG|upright|caption2=Leopard head ornament from the Court of Benin}}
Leopards have been featured in art, mythology and folklore of many countries. In [[Greek mythology]], it was a symbol of the god [[Dionysus]], who was depicted wearing leopard skin and using leopards as means of transportation. In one myth, the god was captured by pirates but two leopards rescued him.<ref name=reaktion>{{cite book |author=Morris, D. |year=2014|title=Leopard|publisher=Reaktion Books|pages=23–24, 31–33, 62, 99, 102, 111 |isbn=9781780233185}}</ref> Numerous Roman mosaics from North African sites depict fauna now found only in [[tropical Africa]].<ref>{{cite journal |title=The Decline of North Africa Since the Roman Occupation: Climatic or Human? |last1=Murphey |first1=R. |journal=Annals of the Association of American Geographers |volume= XLI |issue=2 |year=1951 |pages=116–132 |doi=10.1080/00045605109352048 |url=http://www.geo.hunter.cuny.edu/courses/geog347/articles/decline_human.pdf |archive-url=https://web.archive.org/web/20060914061431/http://www.geo.hunter.cuny.edu/courses/geog347/articles/decline_human.pdf |archive-date=2006-09-14 |url-status=live}}</ref> During the [[Benin Empire]], the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king or ''[[Oba (ruler)|oba]]'', since the leopard was considered the king of the forest.<ref name=britishmuseum>{{cite web |publisher=British Museum |place=London |title=Benin: an African kingdom |url=https://www.britishmuseum.org/PDF/british_museum_benin_art.pdf |archive-url=https://web.archive.org/web/20080805130342/http://www.britishmuseum.org/PDF/british_museum_benin_art.pdf |archive-date=2008-08-05 |url-status=live |access-date=2016-03-29}}</ref> The [[Ashanti people]] also used the leopard as a symbol of leadership, and only the king was permitted to have a ceremonial leopard stool. Some African cultures considered the leopard to be a smarter, better hunter than the lion and harder to kill.<ref name=reaktion/>
In [[Rudyard Kipling]]'s "How the Leopard Got His Spots", one of his ''[[Just So Stories]]'', a leopard with no spots in the [[Highveld]] lives with his hunting partner, the Ethiopian. When they set off to the forest, the Ethiopian changed his brown skin, and the leopard painted spots on his skin.<ref>{{cite book |last1=Kipling |first1=R. |title=Just So Stories |date=1902 |publisher=Macmillan |chapter=How the Leopard Got His Spots |chapter-url=https://theshortstory.co.uk/devsitegkl/wp-content/uploads/2015/06/Short-stories-Rudyard-Kipling-How-the-Leopard-got-his-Spots.pdf |access-date=2023-05-14 |archive-date=2023-11-29 |archive-url=https://web.archive.org/web/20231129190512/https://theshortstory.co.uk/devsitegkl/wp-content/uploads/2015/06/Short-stories-Rudyard-Kipling-How-the-Leopard-got-his-Spots.pdf |url-status=live}}</ref> A leopard played an important role in the 1938 Hollywood film ''[[Bringing Up Baby]]''. African chiefs, European queens, Hollywood actors and [[burlesque]] dancers wore coats made of leopard skins.<ref name=reaktion/>
The leopard is a frequently used motif in [[Leopard (heraldry)|heraldry]], most commonly as ''[[Attitude (heraldry)#Passant|passant]]''.<ref>{{cite book |author=Haist, M. |title=The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature |chapter=The Lion, bloodline, and kingship |pages=3–16 |editor=Hassig, D. |publisher=Taylor & Francis |___location=London |year=1999 |isbn=978-0-8153-2952-7 |chapter-url=https://books.google.com/books?id=lx09r7EOL_oC&pg=PA10 |access-date=2021-09-07 |archive-date=2024-05-05 |archive-url=https://web.archive.org/web/20240505150617/https://books.google.com/books?id=lx09r7EOL_oC&pg=PA10#v=onepage&q&f=false |url-status=live }}</ref> The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the [[Lion (heraldry)|heraldic lion]], and the two are often used interchangeably. Naturalistic leopard-like depictions appear on the [[coat of arms of Benin]], [[Coat of arms of Malawi|Malawi]], [[Coat of arms of Somalia|Somalia]], the [[Coat of arms of the Democratic Republic of the Congo|Democratic Republic of the Congo]] and [[Coat of arms of Gabon|Gabon]], the last of which uses a black panther.<ref>{{cite book|title=The International Flag Book in Color |url=https://archive.org/details/internationalfla00pede|url-access=registration|last=Pedersen|first=C. F. |publisher=Morrow |year=1971}}</ref>
===Attacks on people===
{{Main|Leopard attack}}
The [[Leopard of Rudraprayag]] killed more than 125 people; the Panar Leopard was thought to have killed over 400 people. Both were shot by British hunter [[Jim Corbett]].<ref name=Corbett>{{cite book |title=The Temple Tiger, and More Man-eaters of Kumaon |year=1955 |author=Corbett, J. |author-link=Jim Corbett |publisher=Oxford University Press |place=Oxford}}</ref> The ''spotted devil of Gummalapur'' killed about 42 people in Karnataka, India.<ref>{{cite book |author=Anderson, K. |author-link=Kenneth Anderson (writer) |title=Nine Man-Eaters and one Rogue |publisher=George Allen & Unwin |year=1954 |___location=London |chapter=The Spotted Devil of Gummalapur |pages=36–51 |chapter-url=https://archive.org/details/in.ernet.dli.2015.460900/page/n33/mode/2up}}</ref>
===In captivity===
[[Image:Female animal trainer and leopard, c1906.jpg|thumb|upright|Animal trainer with leopard]]
The [[ancient Roman]]s kept leopards in captivity to be slaughtered in [[Venatio|hunts]] as well as [[Damnatio ad bestias|execute]] criminals.<ref name=reaktion/> In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities.<ref name=britishmuseum/> Several leopards were kept in a menagerie originally established by King [[John of England]] at the [[Tower of London]] in the 13th century; around 1235, three of these animals were given to [[Henry III of England|Henry III]] by Holy Roman Emperor [[Frederick II, Holy Roman Emperor|Frederick II]].<ref>{{cite magazine |last=Owen |first=J. |title=Medieval Lion Skulls Reveal Secrets of Tower of London 'Zoo' |magazine=National Geographic Magazine |date=2005 |url=http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html |archive-url=https://web.archive.org/web/20051105025912/http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html |url-status=dead |archive-date=November 5, 2005 |access-date=2007-09-05}}</ref> In modern times, leopards have been [[Animal training|trained]] and [[Tame animal|tamed]] in [[circus]]es.<ref name=reaktion/>
{{clear}}
==See also==
* [[Leopard pattern]]
* [[List of largest cats]]
* [[Panther (legendary creature)]]
==References==
{{Reflist}}
===Further reading===
* {{Cite book |author-link1=Thomas T. Allsen |last=Allsen |first=Thomas T. |editor-last=Mair |editor-first=V. H. |contribution=Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries |title=Contact and Exchange in the Ancient World |year=2007 |publisher=University of Hawai'i Press |___location=Honolulu |isbn=978-0-8248-2884-4 |ref=none}}
* {{Cite journal |last1=DeRuiter |first1=D. J. |last2=Berger |first2=L. R. |author-link2=Lee R. Berger |year=2000 |title=Leopards as taphonomic agents in dolomitic Caves – Implications for bone accumulations in the hominid-bearing deposits of South Africa |journal=Journal of Archaeological Science |volume=27 |issue=8 |pages=665–684 |doi=10.1006/jasc.1999.0470 |bibcode=2000JArSc..27..665D |ref=none}}
* {{cite book |author=Sanei, A. |title= Analysis of leopard (''Panthera pardus'') status in Iran |url=https://docs.google.com/viewer?a=v&pid=explorer&chrome=true&srcid=0B7Vwy9iDuZifNjliMzYzNTQtZWFjYy00ODgzLWJkNWQtZmY4ZTUzOTY1MTc2&hl=en |publisher=Sepehr Publication Center |language= fa |___location=Tehran |year=2007 |isbn=978-964-6123-74-8 |ref=none }}
* {{Cite journal |author1=Sanei, A. |author2=Zakaria, M. |author3=Yusof, E. |author4=Roslan, M. |year=2011 |title=Estimation of leopard population size in a secondary forest within Malaysia's capital agglomeration using unsupervised classification of pugmarks |journal=Tropical Ecology |volume=52 |issue=1 |pages=209–217 |url=http://www.tropecol.com/pdf/open/PDF_52_2/Jour-07-F.pdf |archive-url=https://web.archive.org/web/20111002102927/http://www.tropecol.com/pdf/open/PDF_52_2/Jour-07-F.pdf |archive-date=2011-10-02 |url-status=usurped |ref=none}}
* {{cite journal|last1=Zakaria |first1= M. |last2= Sanei|first2=A. |year=2011 |url=https://docs.google.com/viewer?a=v&pid=explorer&srcid=0BzbFC60TvnMcNGExNDJjNzItZmM1MC00ODkzLTkzZWMtNjQ3OTAyMzlmOTcz |title=Conservation and management prospects of the Persian and Malayan leopards|journal=Asia Life Sciences |volume=Supplement 7|pages=1–5 |ref=none}}
==External links==
{{Commons and category|Panthera pardus|Panthera_pardus}}
* [http://www.catsg.org/index.php?id=110 IUCN/SSC Cat Specialist Group: ''Panthera pardus'' in Africa]{{Dead link|date=August 2025 |bot=InternetArchiveBot |fix-attempted=yes }} and [http://www.catsg.org/index.php?id=557 ''Panthera pardus'' in Asia] {{Webarchive|url=https://web.archive.org/web/20180623113337/http://www.catsg.org/index.php?id=557 |date=2018-06-23 }}
* {{Cite EB1911|wstitle=Leopard|short=x}}
{{Carnivora|Fe.}}
{{Mammals in culture}}
{{Taxonbar|from=Q34706|from2=Q47450956}}
{{Authority control}}
{{DEFAULTSORT:Leopard}}
[[Category:Leopards| ]]
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[[Category:Big cats]]
[[Category:Felids of Africa]]
[[Category:Felids of Asia]]
[[Category:Mammals described in 1758]]
[[Category:National symbols of Benin]]
[[Category:National symbols of Malawi]]
[[Category:National symbols of Somalia]]
[[Category:National symbols of the Democratic Republic of the Congo]]
[[Category:Panthera]]
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