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Leopard
Temporal range: Early Pleistocene – Present
Male African leopard in Maasai Mara National Reserve, Kenya
CITES Appendix I (CITES)[2]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. pardus[1]
Binomial name
Panthera pardus[1]
Subspecies

See text

Present and historical distribution of the leopard[2]

The leopard (Panthera pardus) is one of the five extant species in the genus Panthera. It has a pale yellowish to dark golden fur with dark spots grouped in rosettes. Its body is slender and muscular reaching a length of 92–183 cm (36–72 in) with a 66–102 cm (26–40 in) long tail and a shoulder height of 60–70 cm (24–28 in). Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).

The leopard was first 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. 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 arid and montane areas. It is an opportunistic predator, hunting mostly ungulates and primates. 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 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.

Etymology[edit]

The English name "leopard" comes from Old French leupart or Middle French liepart, that derives from Latin leopardus and ancient Greek λέοπάρδος (leopardos). Leopardos could be a compound of λέων (leōn), meaning 'lion', and πάρδος (pardos), meaning 'spotted'.[3][4][5] The word λέοπάρδος originally referred to a cheetah (Acinonyx jubatus).[6]

"Panther" is another common name, derived from Latin panther and ancient Greek πάνθηρ (pánthēr);[3] The generic name Panthera originates in Latin panthera, a hunting net for catching wild beasts to be used by the Romans in combats.[7] Pardus is the masculine singular form.[8]

Taxonomy[edit]

Map showing approximate distribution of leopard subspecies

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]

The leopard was designated as the type species of Panthera by Joel Asaph Allen in 1902.[12] In 1917, Reginald Innes Pocock also subordinated the tiger (P. tigris), lion (P. leo), and jaguar (P. onca) to Panthera.[13][14]

Living subspecies[edit]

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]

In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:[17]

Subspecies Distribution Image
African leopard (P. p. pardus) (Linnaeus, 1758)[1] It is the most widespread leopard subspecies and is native to most of Sub-Saharan Africa.[2]
Indian leopard (P. p. fusca) (Meyer, 1794)[18] It occurs in the Indian subcontinent, Myanmar and southern Tibet.[17][19] It is listed as Near Threatened.[20]
Javan leopard (P. p. melas) (Cuvier, 1809)[21] It is native to Java in Indonesia and has been assessed as Endangered in 2021.[22]
Arabian leopard (P. p. nimr) (Hemprich and Ehrenberg, 1830)[23] It is the smallest leopard subspecies and considered endemic to the Arabian Peninsula.[24] 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.[25]
P. p. tulliana (Valenciennes, 1856)[26] 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.[27]

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.[28]

Amur leopard (P. p. orientalis) (Schlegel, 1857)[29][30] It is native to the Russian Far East and northern China, but is locally extinct in the Korean peninsula.[2]
Indochinese leopard (P. p. delacouri) Pocock, 1930[31] It occurs in mainland Southeast Asia and southern China, and is listed as Critically Endangered.[32]
Sri Lankan leopard (P. p. kotiya) Deraniyagala, 1956[33] It is native to Sri Lanka and listed as Vulnerable.[34]

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]

Evolution[edit]

Two cladograms proposed for Panthera. The upper cladogram is based on the 2006[36] and 2009[37] studies, while the lower is based on the 2010[38] and 2011[39] 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 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]

Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion.[40] 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.[41] The leopard-lion clade diverged 3.1–1.95 million years ago.[38][39] Additionally, a 2016 study revealed that the mitochondrial genomes of the leopard, lion and snow leopard are more similar to each other than their nuclear genomes, indicating that their ancestors hybridized with the snow leopard at some point in their evolution.[42]

The oldest unambiguous fossils of the leopard are from Eastern Africa, dating to around 2 million years ago.[43]

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]

Hybrids[edit]

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]

Characteristics[edit]

Mounted skeleton
Rosettes of a leopard
A melanistic leopard or black panther

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]

Its white-tipped tail is about 60–100 cm (23.6–39.4 in) long, white underneath and with spots that form incomplete bands toward the end of the tail.[64] The guard hairs protecting the basal hairs are short, 3–4 mm (0.1–0.2 in) in face and head, and increase in length toward the flanks and the belly to about 25–30 mm (1.0–1.2 in). Juveniles have woolly fur that appear to be dark-coloured due to the densely arranged spots.[61][65] Its fur tends to grow longer in colder climates.[66] The leopard's rosettes differ from those of the jaguar, which are darker and with smaller spots inside.[58] The leopard has a diploid chromosome number of 38.[67]

Melanistic leopards are also known as black panthers. Melanism in leopards is caused by a