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Listed as 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. Leopard skins are popular in fashion.
Felis pardus was the scientific name proposed by Carl Linnaeus in 1758.[9]The generic name Panthera was first used by Lorenz Oken in 1816, who included all the known spotted cats into this group.[10]Oken's classification was not widely accepted, and Felis or Leopardus was used as the generic name until the early 20th century.[11]
Following Linnaeus' first description, 27 leopard subspecies were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered valid on the basis of mitochondrial analysis.[15] Later analysis revealed a ninth valid subspecies, the Arabian leopard.[16]
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.[35]
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 6.37 million years ago. Neofelis diverged about 8.66 million years ago from the Panthera lineage. The tiger diverged about 6.55 million years ago, followed by the snow leopard about 4.63 million years ago and the leopard about 4.35 million years ago. The leopard is a sister taxon to a clade within Panthera, consisting of the lion and the jaguar.[36][37]
Leopard-like fossil bones and teeth possibly dating to the Pliocene were excavated in 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 in southern Hungary.[44]Leopards arrived in Eurasia during the late Early to Middle Pleistocene around 1.2[45] to 0.6 million years ago.[43]Four European Pleistocene leopard subspecies were proposed. P. p. begoueni from the beginning of the Early Pleistocene was replaced about 0.6 million years ago by P. p. sickenbergi, which in turn was replaced by P. p. antiqua around 0.3 million years ago.[46] 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.[46][47]
Leopards depicted in cave paintings in Chauvet Cave provide indirect evidence of leopard presence in Europe.[46]Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central Poland.[48][45]Fossil remains were also excavated in the Iberian[49][50] and Italian Peninsula,[51] and in the Balkans.[52][53]Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago.[54] Leopard fossils were also found in Taiwan.[55]
In 1953, a male leopard and a female lion were 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.[56]
The leopard's fur is generally soft and thick, notably softer on the belly than on the back.[57] Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in rosettes. Its underbelly is white and its ringed tail is shorter than its body. Its pupils are round.[58] 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.[59] 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.[60] Rosette patterns are unique in each individual.[61][62] This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as camouflage.[63]
Melanistic leopards are also known as black panthers. Melanism in leopards is caused by a recessive allele and is inherited as a recessive trait.[68][69][70][71]In India, nine pale and white leopards were reported between 1905 and 1967.[72]Leopards exhibiting erythrism were recorded between 1990 and 2015 in South Africa's Madikwe Game Reserve and in Mpumalanga. The cause of this morph known as a "strawberry leopard" or "pink panther" is not well understood.[73]
The leopard has the largest distribution of all wild cats, occurring widely in Africa, the Caucasus and Asia, although populations are fragmented and declining. It is considered to be locally extinct in North Africa.[2] It inhabits foremost savanna and rainforest, and areas where grasslands, woodlands, and riverine forests remain largely undisturbed.[60] In sub-Saharan Africa, it is still numerous and surviving in marginal habitats where other large cats have disappeared. There is considerable potential for human-leopard conflict due to leopards preying on livestock.[80]
In the Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other Panthera species.[2] 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 2,600 m (8,500 ft) in the Shivalik Hills and Gangetic plains, Central India and Eastern Ghats, Western Ghats, the Brahmaputra River basin and hills in Northeast India.[85] Some leopard populations in the country 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.[86]
In Nepal's Kanchenjunga Conservation Area, a melanistic leopard was photographed at an elevation of 4,300 m (14,100 ft) by a camera trap in May 2012.[87] In Sri Lanka, leopards were recorded in Yala National Park and in unprotected forest patches, tea estates, grasslands, home gardens, pine and eucalyptus plantations.[88][89]In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's Karen State.[90] The Northern Tenasserim Forest Complex in southern Myanmar is considered a leopard stronghold. In Thailand, leopards are present in the Western Forest Complex, Kaeng Krachan-Kui Buri, Khlong Saeng-Khao Sok protected area complexes and in Hala Bala Wildlife Sanctuary bordering Malaysia. In Peninsular Malaysia, leopards are present in Belum-Temengor, Taman Negara and Endau-Rompin National Parks.[91]In Laos, leopards were recorded in Nam Et-Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area.[92][93]In Cambodia, leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest.[94][95]In southern China, leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009.[96]
In Java, leopards inhabit dense tropical rainforests and dry deciduous forests at elevations from sea level to 2,540 m (8,330 ft). Outside protected areas, leopards were recorded in mixed agricultural land, secondary forest and production forest between 2008 and 2014.[97]
The leopard is a solitary and 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, snarls, meows, and purrs.[65] The roaring sequence in leopards consists mainly of grunts,[98] also called "sawing", as it resembles the sound of sawing wood. Cubs call their mother with an urr-urr sound.[65]
The whitish spots on the back of its ears are thought to play a role in communication.[99]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.[100][101]
Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.[60]They can run at over 58 km/h (36 mph; 16 m/s), leap over 6 m (20 ft) horizontally, and jump up to 3 m (9.8 ft) vertically.[98]
Males occupy home ranges 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.[108] 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.[65][75]
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