Leopard
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{{redirect|Leopards}}
{{Taxobox
| name = Leopard
| fossil_range = Late [[Pliocene]] or Early [[Pleistocene]] to Recent
| status = LC
| status_system = iucn2.3
| status_ref = <ref name=IUCN>{{IUCN2006|assessors=Cat Specialist Group|year=2002|id=15954|title=Panthera pardus|downloaded=12 May 2006}} Database entry includes justification for why this species is of least concern</ref>
| trend = down
| image =Leopard africa.jpg
| image_width =200 px
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Carnivora]]
| familia = [[Felidae]]
| genus = ''[[Panthera]]''
| species = '''''P. pardus'''''
| binomial = ''Panthera pardus''
| binomial_authority = [[Carolus Linnaeus|Linnaeus]], 1758
| range_map = Leopard distribution.gif
| range_map_width = 240px
}}
The '''leopard''' ({{IPA2|lɛpɚɖ}}; ''Panthera pardus'') is an [[Old World]] [[mammal]] of the [[Felidae]] [[biological family|family]] and the smallest of the four [[big cats]] of the [[genus]] ''[[Panthera]]'', the other three being the [[tiger]], [[lion]] and [[jaguar]]. Once distributed across southern [[Eurasia]] and [[Africa]], from [[Korea]] to [[South Africa]], the leopard's range of distribution has decreased radically over time because of a variety of factors, including human influence, and the leopard now chiefly occurs in sub-Saharan Africa. There are fragmented populations in [[India]], [[Indochina]], [[Malaysia]], and [[China]]. Despite the loss of range and continual declines in population, the cat remains a "[[Least Concern]]" species; its numbers are greater than that of the other ''Panthera'' species, all of which face more acute conservation concerns.
The leopard has relatively short legs and a long body, with a large skull. Physically, it most closely resembles the jaguar, although it is usually smaller and of slighter build. Its fur is marked with [[rosette (zoology)|rosettes]] which lack internal spots, unlike those of the jaguar. Leopards that are [[melanistic]], either completely black or very dark in coloration, are one of the big cats known colloquially as [[black panther]]s.
The species' success in the wild owes in part to its [[generalist and specialist species|opportunistic]] hunting behaviour, its adaptability to a variety of habitats and its ability to move at up to approximately 60 kilometres (37 miles) an hour. The leopard consumes virtually any animal it can hunt down and catch. Its preferred habitat ranges from [[rainforest]] to [[desert]] terrains. Its [[ecology|ecological]] role and status resembles that of the similarly-sized [[cougar]] in the [[Americas]].
==Etymology==
In [[classical antiquity|antiquity]], it was believed that a leopard was a [[Hybrid (biology)|hybrid]] of a lion and a [[black panther|panther]], as is reflected in its name, a [[Greek language|Greek]] compound word derived from λέων ''léon'' ("lion") and πάρδος ''párdos'' ("male panther"), the latter related to [[Sanskrit language|Sanskrit]] पृदाकु ''pṝdāku'' ("snake, tiger, panther").<ref name=Harper>{{cite web | url = http://www.etymonline.com/index.php?term=leopard | title = "Leopard" | work = Online Etymology Dictionary | publisher = Douglas Harper | accessdate = 2007-11-30}}</ref><ref>{{cite book | last=Monier-Williams | first=Sir Monier | title=A Sanskrit-English Dictionary | coauthors=Prof. E. Leumann, Ph.D., Prof. C. Cappeller, Ph.D., ''et al'' | url=http://www.sanskrit-lexicon.uni-koeln.de/mwview/index.php?sfx=jpg | format=JPEG | accessdate=2007-11-30 | pages=p. 647 | chapter=pṝdāku | chapterurl=http://www.sanskrit-lexicon.uni-koeln.de/cgi-bin/serveimg.pl?file=/MWScan/MWScanjpg/mw0647-pRthukIya.jpg }}</ref>
A ''panther'' can be any of several species of large [[felid]]; in [[North America]], the term refers to [[cougar]]s; in [[South America]], [[jaguar]]s; and everywhere else, it refers to leopards. Early [[natural history|naturalists]] distinguished between leopards and panthers not by colour (a common misconception), but by the length of the tail—panthers having longer tails than leopards.
''Felis pardus'' was one of the many species described in [[Carolus Linnaeus|Linnaeus]]'s 18th-century work, ''[[Systema Naturae]]''.<ref>{{la icon}} {{cite book | last=Linnaeus | first=C | authorlink=Carolus Linnaeus | title=Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. | edition=Editio decima, reformata. | publisher=Holmiae. (Laurentii Salvii). | date=1758 | url=http://dz1.gdz-cms.de/index.php?id=img&no_cache=1&IDDOC=265100}}</ref>
The generic component of its modern scientific designation, ''Panthera pardus'', is derived from Latin via Greek πάνθηρ ''pánthēr''. A [[folk etymology]] held that it was a compound of παν ''pan'' ("all") and θηρ ("beast"). However, it is believed instead to derive from an [[Indo-Iranian]] word meaning "whitish-yellow, pale"; in Sanskrit, this word's reflex was पाण्डर ''pāṇḍara'', from which was derived पुण्डरीक ''puṇḍárīka'' ("tiger", among other things), then borrowed into Greek.<ref name=Harper/><ref>{{cite book | last=Monier-Williams | first=Sir Monier | title=A Sanskrit-English Dictionary | coauthors=Prof. E. Leumann, Ph.D., Prof. C. Cappeller, Ph.D., ''et al'' | url=http://www.sanskrit-lexicon.uni-koeln.de/mwview/index.php?sfx=jpg | format=JPEG | accessdate=2007-11-30 | pages=p. 616 | chapter=pāṇḍara | chapterurl=http://www.sanskrit-lexicon.uni-koeln.de/cgi-bin/serveimg.pl?file=/MWScan/MWScanjpg/mw0616-pANinIyamatadarpaNa.jpg }}</ref><ref>{{cite book | last=Monier-Williams | first=Sir Monier | title=A Sanskrit-English Dictionary | coauthors=Prof. E. Leumann, Ph.D., Prof. C. Cappeller, Ph.D., ''et al'' | url=http://www.sanskrit-lexicon.uni-koeln.de/mwview/index.php?sfx=jpg | format=JPEG | accessdate=2007-11-30 | pages=p. 631 | chapter=puṇḍárīka | chapterurl=http://www.sanskrit-lexicon.uni-koeln.de/cgi-bin/serveimg.pl?file=/MWScan/MWScanjpg/mw0631-puMdhvaja.jpg }}</ref>
==Taxonomy==
Like the rest of the feline family, the ''Panthera'' genus has been subject to much alteration and debate and the exact relations between the four species (as well as the [[clouded leopard]] and [[snow leopard]]) have not been effectively resolved. [[DNA]] evidence shows that the lion, tiger, leopard, jaguar, snow leopard, and clouded leopard share a common ancestor nearly 11 million years ago (Ma)—the [[basal (biology)|basal]] divergence amongst the [[Felidae]] family.<ref name=Johnson2006>{{cite journal | author = Johnson, W.E., Eizirik, E., Pecon-Slattery, J., Murphy, W.J., Antunes, A., Teeling, E. & O'Brien, S.J. | year = 2006 | doi = 10.1126/science.1122277 | title = The Late Miocene radiation of modern Felidae: A genetic assessment. | journal = [[Science (journal)|Science]] | volume = 311 | pages = 73–77 | pmid = 16400146}}</ref> The fossil record points to the emergence of ''Panthera'' just two to 3.8 million years ago.<ref name=Johnson2006/><ref name="Turner1987">{{ cite journal | last = Turner | first = A. | authorlink = | coauthors = | year = 1987 | month = | title = New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora) | journal = Annals of the Transvaal Museum | volume = 34 | issue = | pages = 319–347 | issn = 0041-1752 | url = | accessdate = | quote = }}</ref>
In a [[mitochondrial DNA]] study, Yu and Zhang (2005) suggest that the leopard is most closely related to the snow leopard, and go so far as placing the latter as a fifth species of ''Panthera'', ''P. uncia.''<ref name=Yu>{{cite journal | author = Yu L & Zhang YP | year = 2005 | title = Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores | journal = Molecular Phylogenetics and Evolution | volume = 35 | issue = 2 | pages = 483–495 | doi = 10.1016/j.ympev.2005.01.017|url=http://202.203.208.84/SCI/2005SCI/2005/Li%20Yu%EF%BC%882005M.P.E).pdf|format=PDF}}</ref> Canonical works, such as the Mammal Species of the World, continue to list the snow leopard as the only species within its genus, ''Uncia uncia,''<ref name=msw3>{{MSW3 Wozencraft | pages = 548}}</ref> but this could change; Johnson ''et al.'' (2006) support the placement of the snow leopard within ''Panthera.'' They suggest, however, that the snow leopard is most closely aligned with the tiger. The leopard is held to have diverged from the ''Panthera'' lineage subsequent to these two species, but before the lion and jaguar.<ref name=Johnson2006/> Older research has tended to suggest that the leopard is most closely related to the lion and/or the jaguar. As recently as 2001, it was held to have split along with the lion in a phylogenetic analysis of chemical secretions amongst cats.<ref>{{ cite journal | last =Bininda-Emonds | first =Olaf R.P. | coauthors = Decker-Flum, Denise M.; et al.| year =2001 | month = | title = The utility of chemical signals as phylogenetic characters: an example from the Felidae | journal =Biological Journal of the Linnean Society | volume =72 | issue = | pages =1–15 | format =PDF | url =http://www.personal.uni-jena.de/~b6biol2/Publications/FatCats.pdf | accessdate = 2008-06-07 | quote = }}</ref>
''Panthera'' is believed to have emerged in Asia, with ancestors of the leopard and other cats subsequently migrating into Africa.<ref name=Johnson2006/> Fossil evidence of leopard ancestors has been found from 2 to 3.5 Ma. These [[Pleistocene]] specimens resemble primitive jaguars. The modern leopard type is suggested to have evolved in Africa 470 000–825 000 years ago and radiated across Asia 170 000–300 000 years ago.<ref name=Uphyrkina>{{ cite journal | last =Uphyrkina | first =O. | coauthors =Johnson, E.W.; et al. | year =2001 | month =November | title = Phylogenetics, genome diversity and origin of modern leopard, ''Panthera pardus'' | journal =Molecular Ecology | volume =10 | issue =11 | pages = 2617–2633 | url = http://www.blackwell-synergy.com/links/doi/10.1046/j.0962-1083.2001.01350.x?cookieSet=1 | accessdate =2008-08-06 | doi = 10.1046/j.0962-1083.2001.01350.x }}</ref>
===Subspecies===
[[Image:Panther.jpg|thumb|Indian leopard]]
As many as 27 leopard subspecies were once suggested, the number growing from the time of [[Linnaeus]] in the 18th century to that of [[Reginald Pocock]] in the early 20th. In 1996, Miththapala ''et al.'' revised this downward to just eight subsepecies based on DNA analysis.<ref name=Miththapala>{{ cite journal | last = Miththapala | first =Sriyanie | coauthors =Seidensticker, John; O'Brien, Stephen J. | year =1996 | month =August | title = Phylogeographic Subspecies Recognition in Leopards (''P. pardus''): Molecular Genetic Variation. | journal =Conservation Biology | volume =10 | issue =4 | pages =1115–1132 | id = | url =http://www.blackwell-synergy.com/links/doi/10.1046/j.1523-1739.1996.10041115.x | accessdate = 2008-06-06 | quote = | doi = 10.1046/j.1523-1739.1996.10041115.x }}</ref> Uphyrina ''et al.'' would concur in 2001 but split out a ninth separately, the [[Arabian leopard]] (''P. pardus nimr''). The latter researchers note the number might be an underestimation because of limited sampling of African leopards. Their list as follows:<ref name=Uphyrkina/>
*[[Indo-Chinese leopard]] (''P. pardus delacouri'') in [[Indochina|Mainland Southeast Asia]].
*[[Indian leopard]] (''P. pardus fusca'') in [[India]], southeastern [[Nepal]], and northern [[Bangladesh]].
*[[North Chinese leopard]] (''P. pardus japonensis'') in [[China]].
*[[Sri Lankan leopard]] (''P. pardus kotiya'') in [[Sri Lanka]].
*[[Javan leopard]] (''P. pardus melas'') in [[Java]].
*[[Amur leopard]] (''P. pardus orientalis'') in the Russian [[Far East]], northern China, and [[Korea]].
*[[African leopard]] (''P. pardus pardus'') in Africa.
*[[Persian leopard]] or Iranian leopard (''P. pardus saxicolor'') in [[Southwest Asia]].
*[[Arabian leopard]] (''P. pardus nimr'') in [[Arabian Peninsula]].
[[Image:Leo4.JPG|thumb|Sri Lankan leopard]]
[[Image:leopard walking.jpg|thumb|upright|Female leopard in the Sabi Sands area of South Africa. Note the white spot on its tail, used for communicating with cubs while hunting or in long grass]]
====Older taxonomic divisions====
Included in the African leopard (''P. pardus pardus''):<ref name=Miththapala/>
* [[Barbary leopard]] (''P. pardus panthera'')
* Cape leopard (''P. pardus melanotica'')
* Central African leopard (''P. pardus shortridgei'')
* Congo leopard (''P. pardus ituriensis'')
* East African leopard (''P. pardus suahelica'')
* Eritrean leopard (''P. pardus antinorii'')
* Somalian leopard (''P. pardus nanopardus'')
* Ugandan leopard ((''P. pardus chui'')
* West African leopard (''P. pardus reichinowi'')
* West African forest leopard (''P. pardus leopardus'')
* [[Zanzibar leopard]] (''P. pardus adersi'')
Included in the Persian leopard (''P. pardus saxicolor''):<ref name=Miththapala/>
* [[Anatolian leopard]] (''P. pardus tulliana'')
* Baluchistan leopard (''P. pardus sindica'')
* Caucasus leopard (''P. pardus ciscaucasica'')
* Central Persian leopard (''P. pardus dathei'')
* [[Sinai leopard]] (''P. pardus jarvisi'')
Included in the Indian leopard (''P. pardus fusca''):<ref name=Miththapala/>
* Kashmir leopard (''P. pardus millardi'')
* Nepal leopard (''P. pardus pernigra'')
==Physical characteristics==
The leopard is an agile and stealthy predator. Although smaller than the other members of the ''Panthera'' genus, the leopard is still able to take large prey given a massive skull that well utilizes powerful jaw muscles.<ref name=CAP/> Its body is comparatively long for a cat and its legs are short.<ref name=AWF/> Head and body length is between 90 and 190 cm (35 and 75 in), the tail reaches 60 to 110 cm (24 to 43 in). Shoulder height is 45 to 80 cm (18-31 in). Males are considerably larger than females and weigh 37 to 90 kg (81 to 198 lbs) compared to 28 to 60 kg (62 to 132 lbs) for females.<ref name="Walker's">Ronald M. Nowak: ''Walker's Mammals of the World''. Johns Hopkins University Press, 1999 ISBN 0-8018-5789-9</ref>
One of many spotted cats, a leopard may be mistaken for a cheetah or a jaguar. The leopard has [[rosette (zoology)|rosettes]] rather than [[cheetah]]'s simple spots, but they lack internal spots, unlike the jaguar. The leopard is larger and less lanky than the cheetah but smaller than the jaguar. The leopard's black, irregular rosettes serve as camouflage. They are circular in East Africa but tend to be square-shaped in southern Africa.<ref name=AWF>{{cite web |url=http://www.awf.org/content/wildlife/detail/leopard |title=Leopard |accessdate=2007-09-21 |publisher=[[African Wildlife Foundation]]}}.</ref>
Leopards have been reported to reach 21 years of age in captivity.<ref>{{cite book |last=Crandall |first=L |title=The management of wild animals in captivity |year=1964 |publisher=University of Chicago Press |location=Chicago |isbn= }}</ref>
=== Black leopards ===
{{Main|Black panther}}
A [[melanistic]] morph of the leopard occurs, particularly in mountainous areas and [[rain forests]]. The black color is heritable and caused by [[recessive]] [[gene]] loci.<ref name=Eizirik>{{ cite journal | last =Eizirik | first =Eduardo | coauthors =Yuhki, Naoya; et al. | year = 2003| month =May | title =Molecular genetics and evolution of melanism in the cat family | journal = Current Biology| volume =13 | issue =5 | pages =448–453 | url =http://cat.inist.fr/?aModele=afficheN&cpsidt=14600002 | accessdate = 2008-06-07 | doi =10.1016/S0960-9822(03)00128-3}}</ref> (While they are commonly called [[Black Panther|black panthers]], the term is not exclusive to leopards; it also applies to melanistic jaguars.)
Melanistic leopards are particularly common on the [[Malayan Peninsula]]: early reports suggested up to half of all leopards there are black, but a 2007 camera-trap study in [[Taman Negara National Park]] found that ''all'' specimens were melanistic.<ref name=Malaysia>{{ cite journal | last =Sunquist | first =Fiona | year =2007 | month =Dec/Jan | title = Malaysian Mystery Leopards | journal =National Wildlife Magazine | volume =45 | issue = 1 | url = http://www.nwf.org/nationalwildlife/printerFriendly.cfm?issueID=112&articleID=1413 | accessdate = 2008-06-07 }}</ref> Although the benefits of melanism are difficult to interpret, it may serve as camouflage in the [[rainforest]] habitat. (Genetic research has found four [[convergent evolution|independent origins]] for melanism in cats, suggesting that there must be some adaptive advantage.)<ref name=Eizirik/> Another possibility is that the color variation was relic adaptation to an epidemic; genes causing melanism can also affect the immune system.<ref name=Malaysia/>
In Africa, black leopards are much less common as melanism is not an adaptive advantage on the savanna: dark coloration provides poor camouflage and makes hunting difficult. Estimates are as low as one in 80 or 100. In the dense forests of the [[Ethiopian Highlands]], however, the black leopard is much more common than in Africa generally; as many as one in five leopards may be melanistic.<ref>{{cite web |url=http://www.aau.edu.et/faculties/sc/BSE/wildlife.html#melanism |title=Melanism in Felines
|accessdate=2008-06-07 |work=Wildlife of Ethiopia|publisher=Biological Society of Ethopia |date=April 2001 }}</ref>
==Biology and behavior==
[[Image:leopard rear view soft.jpg|thumb|left|upright|200px|Female leopard. Note the [[Eyespot (mimicry)|ocelli]] (white spots on the back of the ears) used.]]
The leopard is known for its ability in climbing, and it has been observed resting on tree branches during the day and descending from trees headfirst.<ref name=AWD>{{cite web|url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_pardus.html|title=Leopard biology and behavior|accessdate=6 June|accessyear=2008}}</ref> It is a powerful swimmer, although, not as strong as some other big cats, such as the [[tiger]]. The leopard is also very agile, and can run over sixty kilometres an hour, leap over six metres and jump up to three metres vertically.<ref name="animalbytes">{{cite web|url=http://www.seaworld.org/animal-info/Animal-bytes/animalia/eumetazoa/coelomates/deuterostomes/chordata/craniata/mammalia/carnivora/leopard.htm|title=Animal bytes - ''Panthera pardus''|accessdate=6 June|accessyear=2008}}</ref> The leopard is primarily a [[nocturnal]] creature, and many of its operations are done by night. However, there have been recorded instances of leopards hunting during the light, especially when the sky is overcast. It spends much of its day resting and sleeping, up in the branches of tress, underneath rocks or in the grass.<ref name="animalbytes" />
===Diet and hunting===
Leopards are opportunistic hunters. Although mid-sized animals are preferred, the leopard will eat anything from [[dung beetles]] to {{convert|900|kg|lb|0|abbr=on}} male [[giant eland]]s.<ref name=CAP> Nowell, K.; Jackson, P. eds. (1996). [http://carnivoractionplans1.free.fr/wildcats.pdf ''Wild Cats. Status Survey and Conservation Action Plan.''] IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland. (see ''Panthera Pardus'', pp. 24 – 29.)</ref> Their diet consists mostly of [[ungulate]]s and [[monkey]]s, but [[rodent]]s, [[reptile]]s, [[amphibian]]s, [[bird]]s and [[fish]] are also eaten;<ref name ="Schaller290">Schaller, p. 290</ref> In Africa, mid-sized antelopes provide a majority of the leopard's prey, especially [[impala]] and [[Thomson's gazelle]]s.<ref name ="Schaller291">Schaller, p. 291</ref> In Asia the leopard preys on deer such as [[chital]]s and [[muntjac]]s as well as various Asian antelopes and [[Ibex]]. One study at the [[Wolong Reserve]] in China revealed how adaptable the leopard's hunting behaviour is: over the course of seven years vegetative cover receded, and the animals opportunistically shifted from primarily consuming [[tufted deer]] to instead pursuing [[bamboo rat]]s and other smaller prey.<ref>{{ cite journal | last = Johnson| first =Kenneth G. | coauthors =Weng, Wei; et al. | year =1993 | month =August | title =Food Habits of Asiatic Leopards (Panthera pardus fusea) in Wolong Reserve, Sichuan, China | journal =Journal of Mammalogy | volume = 74| issue =3 | pages =646–650 | url =http://www.jstor.org/stable/1382285 | accessdate = 2008-06-13 | quote = | doi =10.2307/1382285 }}</ref>
The leopard stalks its prey silently and at the last minute pounces on its prey and strangles its throat with a quick bite. Leopards often hide their kills in dense vegetation or take them up trees,<ref name ="Schaller291"/> and are capable of carrying animals up to three times their own weight this way.
One survey of nearly 30 research papers found preferred prey weights of 10 to 40 [[kg]] (22-88 lb), with {{convert|25|kg|lb|0|abbr=on}} most preferred. Along with impala and chital, a preference for [[bushbuck]] and [[common duiker]] was found. Other prey selection factors include a preference for prey in small herds, in dense habitat, and those that afford the predator a low risk of injury.<ref>{{cite journal |last=Hayward |first=M. W. |coauthors=Henschel, P.; et al. |year=2006 |month=October |title=Prey preferences of the leopard (Panthera pardus) |journal=Journal of Zoology |volume=270 |issue=2 |pages=298–313 |url=http://www.blackwell-synergy.com/doi/abs/10.1111/j.1469-7998.2006.00139.x |accessdate= 2007-09-13 |doi=10.1111/j.1469-7998.2006.00139.x}}</ref>
[[Image:Leopard on the tree.jpg|thumb|right|200px|[[Anatolian leopard]] resting on a tree]]
===Reproduction===
A male may follow a female who catches his attention. Eventually, a fight for reproductive rights may take place. Depending on the region, leopards may mate all year round (Asia and Africa) or seasonally during January to February (Manchuria and Siberia). The [[estrous cycle]] lasts about 46 days and the female usually is in heat for 6–7 days.<ref>
{{cite book |author=Sadleir R |year=1966 |chapter=Notes on the Reproduction of the larger Felidae |title= Int. Zoo Yearbook: Vol 6|pages=184-87 |publisher=Zool. Soc. London |location=London |isbn= }}</ref> Cubs are usually born in a litter of 2–4,<ref>{{cite web|url=http://www.wildcatconservation.org/Leopard_(Panthera_pardus).html|title=Leopard|publisher=wildcatconservation.org|accessdate=6 June|year=2008}}</ref> but infant mortality is high and usually no more than 1–2 cubs survive beyond their infancy. The pregnant females find a cave, crevice among boulders, hollow tree, or thicket to give birth and make a den. Cubs open their eyes after a period of 10 days. The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in color with less defined spots. Around three months the infants begin to follow the mother out on hunts. At one year of age leopard young can probably fend for themselves but they remain with the mother for 18–24 months.<ref>{{cite web|url=http://brainmuseum.org/Specimens/carnivora/leopard/index.html|title=Leopard (''Pantehra pardus''); Physical characteristics and distribution|accessdate=7 June|accessyear=2008}}</ref>
===Social structure and home range===
Studies of leopard home range size have tended to focus on protected areas, which may have led to skewed data; as of the mid-1980s, only 13% of the leopard range actually fell within a protected area.<ref name=Namibia>{{cite journal |url=http://www.cheetah.org/ama/orig/leopard.pdf |last=Marker |first=L. L. |coauthors=Dickman, A. J.|year=2005 |month=March|title=Factors affecting leopard (''Panthera pardus'') spatial ecology, with particular reference to Namibian farmlands |journal=South African Journal of Wildlife Research |volume=35 |issue=2 |accessdate= 2007-09-13}}</ref> In their IUCN survey of the literature, Nowell and Jackson suggest male home territories vary between 30-78 [[square kilometer]]s, but just 15-16 km² for females.<ref name=CAP/> Research in a conservation area in Kenya shows similar territory sizes and sex differential: 32.8 km² ranges for males, on average, and 14 km² for females.<ref>{{cite journal |last=Mizutani |first=F. |coauthors= Jewell, P. A.|year=1998 |title=Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=40975 |journal=Journal of Zoology|pages=269–286 |volume=244 |doi=10.1017/S0952836998002118|accessdate= 2007-09-13}}</ref> In [[Nepal]], somewhat larger male ranges have been found at about 48 km², while female ranges are in-keeping with other research, at 17 km²; female home territories were seen to decrease to just five to seven km² when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase.<ref> {{cite journal |last= Odden|first= Morten |coauthors=Wegge, Per |year=2005 |title=Spacing and activity patterns of leopards ''Panthera pardus'' in the Royal Bardia National Park, Nepal |journal= Wildlife Biology|volume=11 |pages=145–152 |doi=10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2 |url=http://www.bioone.org/perlserv/?request=get-document&doi=10.2981%2F0909-6396(2005)11%5B145%3ASAAPOL%5D2.0.CO%3B2 |accessdate= 2007-09-13}}</ref> However, significant variations in size of home territories have been suggested across the leopard's range. In Namibia, for instance, research that focussed on spatial ecology in farmlands outside of protected areas found ranges that were consistently above 100 km², with some more than 300 km²; admitting that their data were at odds with others', the researchers also suggested little or no sexual variation in the size of territories.<ref name=Namibia/> Virtually all sources suggest that males do have larger ranges. There seems to be little or no overlap in territory amongst males, although overlap exists between the sexes; one radio-collar analysis in the [[Ivory Coast]] found a female home range completely enclosed within a male's.<ref name=Ivory> {{cite journal |last= Jenny|first= D.|month=November|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 |url=http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=4026878&q=Panthera+pardus+habitat&uid=1040057&setcookie=yes |accessdate= 2007-09-13}}</ref>
The leopard is solitary and, aside from mating, interactions between individuals appear to be infrequent.<ref name=Ivory/> Aggressive encounters have been observed, however. Two of five males studied over a period of a year at a game reserve in South Africa died, both violently. One was initially wounded in a male-male territorial battle over a carcass; taken in by researchers, it was released after a successful convalescence only to be killed by a different male a few months later. A second was killed by another predator, possibly a [[spotted hyena]]. A third of the five was badly wounded in intraspecific fighting, but recovered.<ref>{{cite journal |last=Hunter |first=Luke |coauthors=Balme, Guy; et al. |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 |id= |url=http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf |accessdate= 2007-09-16 |quote= }}</ref>
==Ecology==
===Distribution and habitat===
Data from 1996 found that the leopard has the largest distribution of any wild cat,<ref name=CAP/> occurring most in certain parts of southern Asia and widely in eastern and central Africa,<ref name="kal">{{citeweb|url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_pardus.html|title=Leopard distribution|author=LeeAnn Bies|accessdate=5 June|accessyear=2008}}</ref> although populations before and since have shown a declining trend and are fragmented outside of subsaharan Africa. The IUCN notes that within sub-Saharan Africa the species is "still numerous and even thriving in marginal habitats" where other large cats have disappeared, but that populations in North Africa may be extinct. In Asia, data on distribution is not consistent: populations in Southwest and Central Asia are small and fragmented; in the northeast portion of the range, they are critically endangered; but in the Indian subcontinent, Southeast Asia, and China, the leopard is still relatively abundant.<ref name=IUCN/>
Leopards live mainly in grasslands, woodlands and riverside forests. The animal has primarily been studied in open savannah habitats, which may have biased common descriptions. It is generally considered [[nocturnal]], for instance, but radio-tracking and scat analysis in West Africa has found that rainforest leopards are more likely to be [[diurnal]] and [[crepuscular]]. Forest leopards are also more specialized in prey selection and exhibit seasonal differences in activity patterns.<ref>{{ cite journal | last = Jenny| first =David| coauthors =Zuberbühler, Klaus | year =2005 | month =September | title =Hunting behaviour in West African forest leopards | journal =African Journal of Ecology | volume = 43| issue = 3| pages =197–200 | url =http://www.blackwell-synergy.com/doi/abs/10.1111/j.1365-2028.2005.00565.x | accessdate = 2008-05-23 | doi =10.1111/j.1365-2028.2005.00565.x}}</ref> While associated with the savanna and rainforest, the leopard is exceptionally adaptable: in the Russian Far East, the animal inhabits temperate forests where winter temperatures reach a low of –25 °C.<ref name=Uphyrkina/>
===Ecological role===
Because of their wide habitat range, leopards must compete for food and safety with other large predators such as [[lion]]s, [[tiger]]s, [[spotted hyena]]s and wild [[Canid|dog]]s. These competitors sometimes may steal the leopard's kill or devour its young. A single lion or tiger is capable of killing an adult leopard. Leopards have adapted to live alongside these other predators by hunting at different times of the day, and by avoiding areas frequented by them. In search of safety, the leopard will often stash its young or a recent kill high up in a tree. Lions are occasionally successful in climbing trees and fetching leopard kills,<ref name ="Schaller293">Schaller, p. 293</ref> and if motivated, an adult tiger might also scale a tree to acquire food.
Nowell and Jackson note that resource portioning occurs where the leopard shares range with the lion or tiger: the leopard tends to take smaller prey (usually less than 75 kg) where its large feline cousins are present.<ref name=CAP/> One tropical forest study suggests that leopards do not always avoid the larger cats by hunting at different times. With relatively abundant prey, tigers and leopards were seen to successfully coexist without competitive exclusion or inter-species dominance hierarchies that may be more common to the savanna.<ref>{{ cite journal | last =Karanth | first =K. Ullas | coauthors = Sunquist, Melvin E. | 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 | url =http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=40765 | accessdate = 2008-06-05| doi = 10.1111/j.1469-7998.2000.tb01076.x}}</ref>
==Variant coloration==
[[Image:Blackleopard.JPG|thumb|200px|A melanistic leopard, or "black panther".]]
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A pseudo-melanistic leopard has a normal background colour, but its excessive markings have coalesced so that its back seems to be entirely black. In some specimens, the area of solid black extends down the flanks and limbs; only a few lateral streaks of golden-brown indicate the presence of normal background colour. Any spots on the flanks and limbs that have not merged into the mass of swirls and stripes are unusually small and discrete, rather than forming rosettes. The face and underparts are paler and dappled like those of ordinary spotted leopards. These melanistic leopards are often incorrectly referred to by the general population as "black panthers".<ref>{{cite book|last=Gamble|first=Cyndi|coauthors=Rodney Griffiths|title=Leopards: Natural History & Conservation|year=2004|publisher=Voyageur Press|isbn=0896586561}]</ref>
In a paper about panthers and ounces of Asia, [[Reginald Innes Pocock]] used a photo of a leopard skin from southern India; it had large black-rimmed blotches, each containing a number of dots and it resembled the pattern of a [[jaguar]] or [[clouded leopard]]. Another of Pocock's leopard skins from southern India had the normal rosettes broken up and fused and so much additional pigment that the animal looked like a black leopard streaked and speckled with yellow.
Most other colour morphs of leopards are known only from paintings or museum specimens. There have been very rare examples where the spots of a normal black leopard have coalesced to give a jet black leopard with no visible markings. Pseudo-melanism (abundism) occurs in leopards. The spots are more densely packed than normal and merge to largely obscure the background colour. They may form swirls and, in some places, solid black areas. Unlike a true black leopard the tawny background colour is visible in places. One pseudo-melanistic leopard had a tawny orange coat with coalescing rosettes and spots, but white belly with normal black spots (like a black-and-tan dog).
A 1910 description of a pseudo-melanistic leopard:
{{quote|There is, however, a peculiar dark phase in South Africa, a specimen of which was obtained in 1885 in hilly land covered with scrub-jungle, near Grahamstown. The ground-colour of this animal was a rich tawny, with an orange tinge; but the spots, instead of being of the usual rosette-like form, were nearly all small and solid, like those on the head of an ordinary leopard; while from the top of the head to near the root of the tail the spots became almost confluent, producing the appearance of a broad streak of black running down the back. A second skin had the black area embracing nearly the whole of the back and flanks, without showing any trace of the spots, while in those portions of the skin where the latter remained they were of the same form as in the first specimen. Two other specimens are known; the whole four having been obtained from the Albany district. These dark-coloured South African leopards differ from the black leopards of the northern and eastern parts of Africa and Asia in that while in the latter the rosette-like spots are always retained and clearly visible, in the former the rosettes are lost – as, indeed, is to a considerable extent often the case in ordinary African leopards – and all trace of spots disappears from the blacker portions of the skin.|Lydekker, R. (1910)|''Harmsworth Natural History''}}
Another pseudo-melanistic leopard skin was described in 1915 by Holdridge Ozro Collins who had purchased it in 1912. It had been killed in Malabar, India that same year.
{{quote|The wide black portion, which glistens like the sheen of silk velvet, extends from the top of the head to the extremity of the tail entirely free from any white or tawny hairs … In the tiger, the stripes are black, of a uniform character, upon a tawny background, and they run in parallel lines from the centre of the back to the belly. In this skin, the stripes are almost golden yellow, without the uniformity and parallelism of the tiger characteristics, and they extend along the sides in labyrinthine graceful curls and circles, several inches below the wide shimmering black continuous course of the back. The extreme edges around the legs and belly are white and spotted like the skin of a leopard … The skin is larger than that of a leopard but smaller than that of a full grown tiger.|Collins, Holdridge Ozro (1915)}}
In May 1936, the [[British Natural History Museum]] exhibited the mounted skin of an unusual Somali leopard. The pelt was richly decorated with an intricate pattern of swirling stripes, blotches, curls and fine-line traceries. This is different from a spotted leopard, but similar to a [[king cheetah]] hence the modern [[cryptozoology]] term king leopard. Between 1885 and 1934, six pseudo-melanistic leopards were recorded in the Albany and Grahamstown districts of South Africa. This indicated a mutation in the local leopard population. Other king leopards have been recorded from Malabar in southwestern India. Shooting for trophies may have wiped out these populations.
==Hybrids==
{{Main|Pumapard}}
[[Image:Pumapard-1904.jpg|thumb|Pumapard, taken in 1904.]]
A pumapard is a [[hybrid (biology)|hybrid]] animal resulting from a union between a leopard and a [[puma]]. Three sets of these hybrids were bred in the late 1890s and early 1900s by [[Carl Hagenbeck]] at his animal park in [[Hamburg]], Germany. Most did not reach adulthood. One of these was purchased in 1898 by [[Berlin]] Zoo. A similar hybrid in Berlin Zoo purchased from Hagenbeck was a cross between a male leopard and a female puma. Hamburg Zoo's specimen was the reverse pairing, the one in the black and white photo, fathered by a puma bred to an Indian leopardess.<ref name="pumapard">{{cite web|url=http://media.www.thestrand.ca/media/storage/paper404/news/2005/03/17/Features/Le.Pumapard-891758.shtml|title=Le Pumapard|author=Jen Stevenson|publisher=The Strand|accessdate=16 June|accessyear=2008}}</ref>
Whether born to a female puma mated to a male leopard, or to a male puma mated to a female leopard, pumapards inherit a form of dwarfism. Those reported grew to only half the size of the parents. They have a puma-like long body (proportional to the limbs, but nevertheless shorter than either parent), but short legs. The coat is variously described as sandy, tawny or greyish with brown, chestnut or "faded" rosettes.<ref name="pumapard" />
==Leopards and humans==
[[Image:Dionysus on a leopard, pella, greece.jpg|200px|thumb|left|A [[mosaic]] depicting [[Dionysus]] on a leopard. 4th century BC. [[Pella]], [[Greece]].]]
Leopards have been known to humans since antiquity and have featured in the art, mythology and folklore of many countries where they have historically occurred, such as [[ancient Greece]], [[Persia]] and Rome, as well as some where they have not existed since for several millenniums, such as in [[England]]. The modern use of the leopard as an emblem for sport or a coat of arms is much more restricted to Africa, though numerous products worldwide have used the name.
Leopards and humans have many relations, involving [[tourism]], [[heraldry]] and modern culture. Leopard domestication has also been recorded - several leopards were kept in a menagerie established by [[John of England|King John]] at the [[Tower of London]] in the 13th century; around 1235 of these animals were given to [[Henry III of England|Henry III]] by the Holy Roman Emperor [[Frederick II, Holy Roman Emperor|Frederick II]].<ref>{{cite web | last = Owen | first = James | title = Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo" | work = National Geographic Magazine | publisher = National Geographic | date = November 3, 2005 | url = http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html | accessdate = 2007-09-05}}</ref>
[[Image:Leopard near driver.jpg|thumb|right|200px|A female leopard in the Sabi Sands of South Africa illustrating just how close tourists can get to these wild cats.]]
===Tourism===
Despite its size, this largely [[nocturnal]] and [[arboreal]] [[predator]] is difficult to see in the wild. The best location to see leopards in Africa is in the Sabi Sand Private Game Reserve in [[South Africa]], where leopards are habituated to [[safari]] vehicles and are seen on a daily basis at very close range. In Asia, one can see leopards at [[Yala National Park]] in [[Sri Lanka]], which has one of the world's highest densities of wild leopards, but even here sightings are by no means guaranteed because more than half the park is closed to the public, allowing the animals to thrive. Another good destination for leopard watching is the recently reopened [[Wilpattu National Park]], also in Sri Lanka. In [[India]] leopards are found all over the country and this wide distribution leads to maximum man-animal conflict. Among the best places to observe leopards in India are national parks in [[Madhya Pradesh]] and [[Uttarakhand]].
[[Image:Grosses Landeswappen Baden-Wuerttemberg.png|left|180px|thumb|Coat of arms of the German state of Baden-Württemberg]]
===Heraldry===
{{main|Leopard (heraldry)}}
The lion passant guardant or "leopard" is a frequently used [[charge (heraldry)|charge]] in [[heraldry]], most commonly appearing in groups of three.<ref>{{cite book|title=The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature|last=Strickland|first=Debra Higgs|coauthors=Debra Hassig|publisher=Taylor & Francis|year=1999|isbn=0815329520}}</ref> The heraldric leopard lacks spots and sports a mane, making it visually almost identical to the [[lion (heraldry)|heraldric lion]], and the two are often used interchangeably. These traditional lion passant guardants appear in the [[coat of arms of England]] and many of its former colonies; more modern naturalistic (leopard-like) depictions appear on the coat of arms of several African nations including [[coat of arms of Benin|Benin]], [[Coat of arms of Malawi|Malawi]], [[Coat of arms of Somalia|Somalia]], [[Coat of arms of the Democratic Republic of the Congo|Democratic Republic of the Congo]] and [[Coat of arms of Gabon|Gabon]] which uses a black panther.<ref>{{cite book|title=The International Flag Book in Color|last=Pedersen|first=Christian Fagd|publisher=Morrow|year=1971}}</ref>
===Man-eating===
Although most leopards will tend to avoid humans, people are occasionally targeted as prey. Most healthy leopards prefer wild prey to humans, but cats who are injured, sickly or struggling with a shortage of regular prey often turn to hunting people and may become habituated to it. In the most extreme cases, both in India, a leopard dubbed "the [[Leopard of Rudraprayag]]" is claimed to have killed over 125 people and the infamous leopard called "[[Panar Leopard]]" killed over 400 after being injured by a poacher and thus being made unable to hunt normal prey.<ref>{{cite book|title=Man the Hunted: Primates, Predators, and Human Evolution|last=Hart|first=Donna|coauthors=Robert W. Sussman|publisher=Westview Press|year=2005|isbn=0813339367}}</ref><ref>Tougias, p.147</ref> The "Leopard of Rudraprayag" and the "Panar Leopard" were both killed by the famed hunter [[Jim Corbett (hunter)|Jim Corbett]].<ref>{{cite book|title=Death in the Long Grass|last=Capstick|first=Peter Hathaway|publisher=St. Martin's Press|year=1978|isbn=0312186134}}</ref> Man-eating leopards are considered bold by feline standards and commonly enter human settlements for prey, more so than their lion and tiger counterparts.<ref>The Spotted Devil of Gummalapur, ''Nine Man-Eaters and one Rogue'', Kenneth Anderson, Allen & Unwin Ltd, 1954</ref> [[Kenneth Anderson (writer)|Kenneth Anderson]], who had first hand experience with many man-eating leopards, described them as far more threatening than tigers:
[[Image:Spotted-Devil.jpg|thumb|The body of the [[Leopard of Gummalapur]], responsible for the deaths of 42 people]]
{{bquote|Although examples of such animals are comparatively rare, when they do occur they depict the panther [leopard] as an engine of destruction quite equal to his far larger cousin, the tiger. Because of his smaller size he can conceal himself in places impossible to a tiger, his need for water is far less, and in veritable demoniac cunning and daring, coupled with the uncanny sense of self preservation and stealthy disappearance when danger threatens, he has no equal...|x|x|''Nine Man-Eaters and One Rogue'', Chapter II ''The Spotted Devil of Gummalapur''}}
Because they can subsist on small prey and are less dependent on large prey, leopards are less likely to turn to man-eating than either lions or tigers. However, leopards might be attracted to human settlements by livestock or pets, especially domestic dogs, and they may resort to the eating of humans should conditions demand it, and no other food is available.<ref>{{cite web|url=http://209.85.173.104/search?q=cache:PsxflauUcl0J:3w.terrasco.net/files/Nature/Felidae/Panthera/Panthera_pardus.pdf+panthera+pardus+man-eating&hl=en&ct=clnk&cd=7&gl=au&client=firefox-a|title=Panthera pardus: The Leopard|accessdate=8 June|accessyear=2008|author=Markus Fumagalli}}</ref> In addition, leopards also preyed on man's hominid ancestors.<ref>[http://www.cartage.org.lb/en/themes/Sciences/LifeScience/PhysicalAnthropology/HumanGeneticEvolution/EarlyHominids/EarlyHominids.htm Discovery of Early Hominids]</ref>
==See also==
* [[Leopard men]]
==Notes==
{{reflist|2}}
==References==
* Allsen, Thomas T. (2006). "Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries." In: ''Contact and Exchange in the Ancient World''. Ed. Victor H. Mair. University of Hawai'i Press. Pp. 116-135. ISBN-13: ISBN 978-0-8248-2884-4; ISBN-10: ISBN 0-8248-2884-4
* Khalaf-von Jaffa, Norman Ali Bassam Ali Taher (2005). [http://www.geocities.com/jaffacity/Arabischer_Leopard.html The Arabian Leopard (Panthera pardus nimr)]. Gazelle: The Palestinian Biological Bulletin. Number 42, June 2005. pp. 1-8. (in German).
* Khalaf-Sakerfalke von Jaffa, Norman Ali Bassam Ali Taher (2006). [http://www.geocities.com/jaffacity/Chinese_Leopard.html The Chinese leopard (Panthera pardus japonensis, Gray 1862) in Neunkirchen Zoo, Neunkirchen, Saarland, Germany]. Gazelle: The Palestinian Biological Bulletin. Number 60, December 2006. pp. 1-10.
*{{cite book |last=Schaller |first=George B. |year=1972 |title= The Serengeti Lion|publisher= University of Chicago Press|location=Chicago |isbn=0-226-73639-3}}
* Leopards and spots on ears and tail [http://www.sandiegozoo.org/animalbytes/t-leopard.html]
* DeRuiter, D.J. and [[Lee R. Berger|Berger, L.R.]] (2000) Leopards as Taphonomic Agents in dolomitic Caves - Implications for bone Accumulations in the Hominid-bearing Deposits of South Africa. J. Arch. Sci. 27, 665-684.
* {{cite book|title=Beyond Conservation: A Wildland Strategy|last=Taylor|first=Peter|coathors=Stephanie Barrientos, Catherine Dolan|publisher=Earthscan|year=2005|isbn=1844071979}}
* {{cite book|title=When Man Is the Prey: True Stories of Animals Attacking Humans|last=Tougias|first=Michael|publisher=Macmillan|year=2007|isbn=0312373007}}
{{refend}}
==External links==
{{portal|Cats|Cat03.jpg}}
{{commonscat|Panthera pardus}}
{{Wikispecies|Panthera pardus}}
* [http://www.theanimalfiles.com/mammals/carnivores/leopard.html Pictures and Information on Leopards]
* [http://leopards.wild-cat.org/ Leopards at wild-cat.org]
* [http://www.landmarkfoundation.org.za/leopard.htm South African Leopard and Predator Conservation]
* [http://www.awf.org/wildlives/147 Leopard: Wildlife summary from the African Wildlife Foundation]
* [http://about-south-africa.com/html/leopard.html African leopard]
* [http://www.nature.org/animals/mammals/animals/leopard.html The Nature Conservatory's Species Profile: Leopard]
* [http://www.persianleopardcs.org/ Persian Leopard Conservation Society]
* [http://www.arkive.org/species/GES/mammals/Panthera_pardus_nimr/ Images and movies of the South Arabian leopard ''(Panthera pardus nimr)''] from ARKive
* [http://www.arkive.org/species/GES/mammals/Panthera_pardus_kotiya/ Images and movies of the Sri Lankan leopard ''(Panthera pardus kotiya)''] from ARKive
*[http://www.bigcatcare.org Center for Animal Research and Education] Providing Sanctuary for over 50 big cats
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