Leopard

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Leopard
Temporal range: Early Pleistocene – Present
Male
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

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

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

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

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

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);

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

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

Following Linnaeus' first description, 27 leopard

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
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
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

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

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

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

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

In 1953, a male leopard and a female lion were

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

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.

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

diploid chromosome number of 38.[67]

Melanistic leopards are also known as

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]

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.[64] Males stand 60–70 cm (24–28 in) at the shoulder, while females are 57–64 cm (22–25 in) tall. The head-and-body length ranges between 92 and 183 cm (36 and 72 in) with a 66 to 102 cm (26 to 40 in) long tail. Sizes vary geographically. Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).[74] Occasionally, large males can grow up to 91 kg (201 lb). Leopards from the Cape Province in South Africa are generally smaller, reaching only 20–45 kg (44–99 lb) in males.[65][66][75]
The maximum recorded weight of a wild leopard in Southern Africa was around 96 kg (212 lb), and it measured 262 cm (103 in).[76] In 2016, an Indian leopard killed in Himachal Pradesh measured 261 cm (103 in) with an estimated weight of 78.5 kg (173 lb); it was perhaps the largest known wild leopard in India.[77][78]

The largest recorded skull of a leopard was found in India in 1920 and measured 28 cm (11 in) in

basal length, 20 cm (7.9 in) in breadth, and weighed 1 kg (2.2 lb). The skull of an African leopard measured 286 mm (11.3 in) in basal length, and 181 mm (7.1 in) in breadth, and weighed 790 g (28 oz).[79]

Distribution and habitat

Leopard in a tree in India

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]

Leopard populations in the Arabian Peninsula are small and fragmented.[81][82][83] In southeastern Egypt, a leopard killed in 2017 was the first sighting of the species in this area in 65 years.[84]

In the

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

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 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 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

agricultural land, secondary forest and production forest between 2008 and 2014.[97]

In the Russian Far East, it inhabits temperate coniferous forests where winter temperatures reach a low of −25 °C (−13 °F).[16]

Behaviour and ecology

The leopard is a solitary and

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 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 1–25 km (0.62–15.53 mi) across their range at night; wandering up to 75 km (47 mi) if disturbed.[65][75] In some regions, they are nocturnal.[102][103] In western African forests, they have been observed to be largely diurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.[104]

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]

  • A female leopard with white markings on the backs of her ears.
    A female leopard with white markings on the backs of her ears.
  • A female leopard showing white spots on the tail
    A female leopard showing white spots on the tail
  • A leopard climbing down a tree
    A leopard climbing down a tree
  • A leopard hunting a bushpig
    A leopard hunting a bushpig
  • Video of a leopard in the wild

Social spacing

scent-marking his territory[105]

In Kruger National Park, most leopards tend to keep 1 km (0.62 mi) apart.[106] Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.[74][107] Aggressive encounters are rare, typically limited to defending territories from intruders.[66] In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.[102]

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]

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.[109] Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the

Bardia National Park, male home ranges of 48 km2 (19 sq mi) and female ones of 5–7 km2 (1.9–2.7 sq mi) are smaller than those generally observed in Africa.[113]

Hunting and diet

The leopard is a

gray langurs. Leopards also kill smaller carnivores like black-backed jackal, bat-eared fox, genet and cheetah.[114]

The largest prey killed by a leopard was reportedly a male eland weighing 900 kg (2,000 lb).[98] 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 bamboo rats and other smaller prey.[115]

The leopard depends mainly on its acute senses of hearing and vision for hunting.[116] It primarily hunts at night in most areas.[65] In western African forests and Tsavo National Park, they have also been observed hunting by day.[117] They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.[118]

It stalks its prey and tries to approach as closely as possible, typically within 5 m (16 ft) 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 by the throat and strangles them.[65] It caches kills up to 2 km (1.2 mi) apart.[74] 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 125 kg (276 lb) up 5.7 m (18 ft 8 in) into a tree.[117] It eats small prey immediately, but drags larger carcasses over several hundred meters 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.[66][119]

Average daily consumption rates of 3.5 kg (7 lb 11 oz) were estimated for males and of 2.8 kg (6 lb 3 oz) for females.

gemsbok cucumbers, watermelon and Kalahari sour grass.[120]

Enemies and competitors

A lioness steals a leopard kill in Kruger National Park

In parts of its range, the leopard is

sympatric with other large predators such as the tiger, lion, cheetah, spotted hyena, striped hyena, brown hyena, African wild dog, dhole, wolf and up to five bear species. Some of these species steal its kills, kill its cubs and even kill adult leopards. Leopards retreat up a tree in the face of direct aggression, and were observed when killing or preying on smaller competitors such as black-backed jackal, African civet, caracal, and African wildcat.[60][121] Leopards generally seem to avoid encounters with adult bears, killing vulnerable bear cubs instead. In Sri Lanka, a few recorded fights between leopards and sloth bears apparently result in both animals winding up either dead or grievously injured.[122][123] Leopards generally avoid large packs of African wild dogs and dholes, and flee up a tree at the sight of them.[98]

While interspecies killing of full-grown leopards is generally rare, given the opportunity, both the tiger and lion readily kill and consume both young and adult leopards.[118][121][124][125] In the Kalahari Desert, leopards frequently lose kills to brown hyenas, if they are unable to move the kill up a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills.[126][127] Lions occasionally fetch leopard kills from trees.[119]

Resource partitioning occurs where leopards share their range with tigers. Leopards tend to kill smaller prey, usually less than 75 kg (165 lb), where tigers are present.[60]
In areas where leopards and tigers are sympatric, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous.[124] Tigers appear to inhabit the deep parts of the forest while leopards are pushed closer to the fringes.[128] In tropical forests, leopards do not always avoid the larger cats by hunting at different times. With relatively abundant prey and differences in the size of the selected prey, tigers and leopards seem to successfully coexist without competitive exclusion or interspecies dominance hierarchies.[129] In Africa, leopards do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around the same time of day. In addition, there is no evidence that lions effect leopard abundance.[130]

Nile crocodiles occasionally prey on leopards. In one occasion, a large adult leopard was grabbed and consumed by a large crocodile while attempting to hunt along a river bank in Kruger National Park.[106][74] Mugger crocodiles reportedly killed an adult leopard in Rajasthan.[131] An adult leopard was recovered from the stomach of a 5.5 m (18 ft 1 in) Burmese python (Python bivittatus).[132] In the Serengeti National Park, troops of around 30–40 olive baboons were observed mobbing and attacking a female leopard and her cubs.[133]

Reproduction and life cycle

Leopards mating
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.[60] The female's estrous cycle lasts about 46 days, and she is usually in heat for 6–7 days.[134] Gestation lasts for 90 to 105 days.[135] Cubs are usually born in a litter of 2–4 cubs.[136] The mortality rate of cubs is estimated at 41–50% during the first year.[106] Lions and spotted hyenas are the biggest cause for leopard cub mortality during their first year. Male leopards are known to cause infanticide, in order to bring the female back into heat.[74] Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.[60]

Females give birth in a cave, crevice among boulders, hollow tree or thicket. Newborn cubs weigh 280–1,000 g (9.9–35.3 oz), and are born with closed eyes, which open four to nine days after birth.[98][66] 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.[74] 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.[137] After separating from their mother, sibling cubs may travel together for months.[60] Both male and female leopards typically reach sexual maturity at 2–2⅓ years.[74]

The generation length of the leopard is 9.3 years.[138] The average life span of a leopard is 12–17 years.[98] The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.[139]

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.[2]
In 2007, a leopard reintroduction programme was initiated in the Russian Caucasus, where captive bred individuals are reared and trained in 0.5–0.9 ha (1.2–2.2 acres) large enclosures in

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.[2] Contemporary records suggest that the leopard occurs in only 25% of its historical range.[141][142]

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.[143]
In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.[144] In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.[145]

Surveys in the Central African Republic's

transhumant pastoralists 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.[146]

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.[147]

Human relations

Cultural significance

Leopards on the Magerius Mosaic from modern Tunisia
Leopard head ornament from the Court of Benin

Leopards have been featured in art, mythology and folklore of many countries. In

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.[148]

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.[151] 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.[148]

The leopard is a frequently used motif in

Democratic Republic of the Congo and Gabon, the last of which uses a black panther.[153]

Attacks on people

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.[154] The spotted devil of Gummalapur killed about 42 people in Karnataka, India.[155]

In captivity

Animal trainer with leopard

The

John of England at the Tower of London in the 13th century; around 1235, three of these animals were given to Henry III by Holy Roman Emperor Frederick II.[156] In modern times, leopards have been trained and tamed in circuses.[148]

See also

References

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Further reading

External links