Present and historical distribution of the leopard[2]
Theleopard (Panthera pardus) is one of the five extant catspecies in thegenusPanthera. It has a pale yellowish to dark golden fur with dark spots grouped inrosettes. 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 is adapted to a variety of habitats ranging fromrainforest tosteppe, includingarid and montane areas. It is anopportunistic predator, hunting mostlyungulates andprimates. It relies on its spotted pattern forcamouflage as it stalks and ambushes its prey, which it sometimes drags up a tree. It is asolitary 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 reachsexual maturity at the age 2–2.5 years.
Listed asVulnerable on theIUCN Red List, leopard populations are currently threatened byhabitat loss and fragmentation, and are declining in large parts of the global range. Leopards have had cultural roles inAncient Greece,West Africa and modern Western culture.Leopard skins are popular in fashion.
Etymology
The English name "leopard" comes fromOld Frenchleupart orMiddle Frenchliepart, that derives fromLatinleopardus andancient Greekλεόπαρδος (leopardos).Leopardos could be acompound ofλέων (leōn), meaning'lion', andπάρδος (pardos), meaning'spotted'.[3][4][5] The wordλεόπαρδος originally referred to acheetah (Acinonyx jubatus).[6]
"Panther" is another common name, derived from Latinpanther and ancient Greekπάνθηρ (pánthēr);[3] Thegeneric namePanthera originates in Latinpanthera, a hunting net for catching wild beasts to be used by theRomans in combats.[7]Pardus is themasculine singular form.[8]
Taxonomy
Map showing approximate distribution of leopard subspecies
Following Linnaeus' first description, 27 leopardsubspecies were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been consideredvalid on the basis ofmitochondrial analysis.[15] Later analysis revealed a ninth valid subspecies, theArabian leopard.[16]
In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:[17]
It is the smallest leopard subspecies and consideredendemic to theArabian Peninsula.[24] As of 2023, the population was estimated to comprise 100–120 individuals inOman andYemen; it was therefore assessed asCritically Endangered in 2023.[25] It is locally extinct in Syria, Lebanon, Israel, Palestine, Jordan, Kuwait and the United Arab Emirates.[2]
The Balochistan leopard population in the south of Iran, Afghanistan and Pakistan is separated from the northern population by theDasht-e Kavir andDasht-e Lut deserts.[28]
Results of ananalysis of molecular variance and pairwisefixation 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 forPanthera. 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 ofphylogenetic studies based onnuclear DNA andmitochondrial DNA analysis showed that the lastcommon ancestor of thePanthera andNeofelis genera is thought to have lived about6.37 million years ago.Neofelisdiverged about8.66 million years ago from thePantheralineage. The tiger diverged about6.55 million years ago, followed by the snow leopard about4.63 million years ago and the leopard about4.35 million years ago. The leopard is asister taxon to aclade withinPanthera, consisting of the lion and the jaguar.[36][37]
Results of a phylogenetic analysis of chemicalsecretions amongst cats indicated that the leopard is closely related to the lion.[40]The geographic origin of thePanthera is most likely northern Central Asia. The leopard-lion clade was distributed in the Asian and AfricanPalearctic since at least the earlyPliocene.[41] The leopard-lion clade diverged 3.1–1.95 million years ago.[38][39] Additionally, a 2016 study revealed that themitochondrial genomes of the leopard, lion andsnow leopard are more similar to each other than theirnuclear genomes, indicating that their ancestorshybridized with the snow leopard at some point in theirevolution.[42]
The oldest unambiguousfossils of the leopard are from Eastern Africa, dating to around 2 million years ago.[43]
Leopards depicted in cave paintings inChauvet Cave provide indirect evidence of leopard presence in Europe.[46]Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-centralPoland.[48][45]Fossil remains were also excavated in the Iberian[49][50] andItalian Peninsula,[51][52] and in theBalkans.[53][54]Leopard fossils dating to the Pleistocene were also excavated in theJapanese archipelago.[55] Leopard fossils were also found inTaiwan.[56]
In 1953, a male leopard and a female lion werecrossbred in Hanshin Park inNishinomiya, Japan. Their offspring known as aleopon 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.[57]
Characteristics
Mounted skeleton
3D model of 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.[58] Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped inrosettes. Its underbelly is white and its ringed tail is shorter than its body. Its pupils are round.[59] Leopards living in arid regions are pale cream, yellowish toochraceous andrufous 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.[60] 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 inrainforest habitats.[61] Rosette patterns are unique in each individual.[62][63] This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves ascamouflage.[64]
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.[65]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.[62][66] Its fur tends to grow longer in colder climates.[67]The leopard's rosettes differ from those of thejaguar, which are darker and with smaller spots inside.[59] The leopard has adiploidchromosome number of 38.[68]
The leopard is a slender and muscular cat, with relatively short limbs and a broad head. It issexually dimorphic with males larger and heavier than females.[65] 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).[75] Occasionally, large males can grow up to 91 kg (201 lb). Leopards from theCape Province in South Africa are generally smaller, reaching only 20–45 kg (44–99 lb) in males.[66][67][76]The heaviest wild leopard in Southern Africa weighed around 96 kg (212 lb), and it measured 262 cm (103 in).[77] In 2016, an Indian leopard killed inHimachal 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.[78]
The largest recorded skull of a leopard was found in India in 1920 and measured 28 cm (11 in) inbasal 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]
The leopard has the largest distribution of all wild cats, occurring widely in Africa and Asia, although populations are fragmented and declining.[2] It inhabits foremostsavanna andrainforest, and areas wheregrasslands,woodlands andriparian forests remain largely undisturbed.[61] It also persists in urban environments, if it is not persecuted, has sufficient prey and patches of vegetation for shelter during the day.[80]
The leopard's range inWest Africa is estimated to have drastically declined by 95%, and in theSahara desert by 97%.[81] Insub-Saharan Africa, it is still numerous and surviving in marginal habitats where other large cats have disappeared.[82] In southeasternEgypt, an individual found killed in 2017 was the first sighting of the leopard in this area in 65 years.[83]
In theIndian subcontinent, the leopard is still relatively abundant, with greater numbers than those of otherPanthera species.[2] Some leopard populations inIndia 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.[90] 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 theShivalik Hills andGangetic plains,Central India andEastern Ghats,Western Ghats, theBrahmaputra River basin and hills inNortheast India.[91]In Nepal'sKanchenjunga Conservation Area, a melanistic leopard was photographed at an elevation of 4,300 m (14,100 ft) by a camera trap in May 2012.[92]
The leopard is a solitary andterritorial 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 cannot obtain any prey. They produce a number of vocalizations, including growls and snarls. Cubs call their mother with meows and anurr-urr sound.[66] The most notable vocalization is the 'sawing'roar, which consists of deep, repeated strokes. This likely functions in establishing territories and attracting mates.[103]
The whitish spots on the back of its ears are thought to play a role in communication.[104]It has been hypothesized that the white tips of their tails may function as a 'follow-me' signal inintraspecific communication. However, no significant association were found between a conspicuous colour of tail patches and behavioural variables in carnivores.[105][106]
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 up to 25 km (16 mi) across their range at night; wandering up to 75 km (47 mi) if disturbed.[66][76][107][108] In western African forests, they have been observed to be largelydiurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.[109]
Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.[61]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.[110]
A female leopard with white markings on the backs of her ears
InKruger National Park, most leopards tend to keep 1 km (0.62 mi) apart.[112] Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.[75][113] Aggressive encounters are rare, typically limited to defending territories from intruders.[67] In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.[107]
Males occupyhome ranges that often overlap with a few smaller female home ranges, probably as a strategy to enhance access to females. In theIvory Coast, the home range of a female was completely enclosed within a male's.[114] 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.[66][76]
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.[115]Sizes of home ranges vary geographically and depending on habitat and availability of prey. In theSerengeti, males have home ranges of 33–38 km2 (13–15 sq mi) and females of 14–16 km2 (5.4–6.2 sq mi);[116][117] but males in northeastern Namibia of 451 km2 (174 sq mi) and females of 188 km2 (73 sq mi).[118] They are even larger in arid and montane areas.[67] In Nepal'sBardia 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.[119]
The leopard is acarnivore that prefers medium-sized prey with a body mass ranging from 10–40 kg (22–88 lb). Prey species in this weight range tend to occur in dense habitat and to form small herds. Species that prefer open areas and have well-developed anti-predator strategies are less preferred. More than 100 prey species have been recorded. The most preferred species areungulates, such asimpala,bushbuck,common duiker andchital.Primates preyed upon includewhite-eyelid mangabeys,guenons andgray langurs. Leopards also kill smaller carnivores likeblack-backed jackal,bat-eared fox,genet and cheetah.[120] In urban environments,domestic dogs provide an important food source.[80] The largest prey killed by a leopard was reportedly a maleeland weighing 900 kg (2,000 lb).[110] A study inWolong 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 consumingtufted deer to pursuingbamboo rats and other smaller prey.[121]
The leopard depends mainly on its acute senses of hearing and vision for hunting.[122] It primarily hunts at night in most areas.[66] In western African forests and Tsavo National Park, they have also been observed hunting by day.[123] They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.[124] 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 animalsby the throat and strangles them.[66] Itcaches kills up to 2 km (1.2 mi) apart.[75] 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.[123] It eats small prey immediately, but drags larger carcasses over several hundred metres and caches it safely in trees, bushes or even caves; this behaviour allows the leopard to store its prey away from rivals, and offers it an advantage over them. The way it stores the kill depends on local topography and individual preferences, varying from trees in Kruger National Park to bushes in the plain terrain of the Kalahari.[67][125] Before their extirpation from Europe, leopards there cached their meat in caves, as evidenced by fossilised bone accumulations in caves such as Los Rincones in theProvince of Zaragoza, Spain.[126]
Leopards are known to drop from trees onto Impalas, which is probably an opportunistic hunting behaviour. A leopard falling from a height of 2.69 metres onto the back of its prey (3.55 metres total height), takes 0.7 seconds to fall and reaches a terminal velocity of 25 km/h; this hunting technique requires that the prey be unaware of the predator's attack and it also requires great precision to avoid falling on the horns of males, which allows for a safe attack.[127]
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.[112] In the southernKalahari Desert, leopards meet their water requirements by the bodily fluids of prey andsucculent plants; they drink water every two to three days and feed infrequently on moisture-rich plants such asgemsbok cucumbers,watermelon and Kalaharisour grass.[128]
Enemies and competitors
A lioness steals a leopard kill in Kruger National Park
Across its range, the leopard coexists with a number of other large predators. In Africa, it is part of a large predatorguild with lions, cheetahs,spotted andbrown hyenas, andAfrican wild dogs.[129] The leopard is dominant only over the cheetah while the others have the advantage of size, pack numbers or both.[66] Lions pose a great mortal threat and can be responsible for 22% of leopard deaths inSabi Sand Game Reserve. Spotted hyenas are less threatening but are more likely tosteal kills, being the culprits of up to 50% of stolen leopard kills in the same area.[130][131] To counter this, leopards store their kills in the trees and out of reach.[131][132] Lions have a high success rate in fetching leopard kills from trees.[131] Leopards do not seem to actively avoid their competitors but rather difference in prey and habitat preferences appear to limit their spatial overlap.[129] In particular, leopards use heavy vegetation regardless of whether lions are present in an area and both cats are active at the same time of day.[133]
In Asia, the leopard's main competitors are tigers anddholes. Both the larger tiger and pack-living dhole dominate leopards during encounters. Interactions between the three predators involve chasing, stealing kills and direct killing.[134] Tigers appear to inhabit the deep parts of the forest while leopards and dholes are pushed closer to the fringes.[135] The three predators coexist by hunting different sized prey.[136] InNagarhole National Park, the average size for a leopard kill was 37.6 kg (83 lb) compared to 91.5 kg (202 lb) for tigers and 43.4 kg (96 lb) for dholes.[137] AtKui Buri National Park, following a reduction in prey numbers, tigers continued to feed on favoured prey while leopards and dholes had to increase their consumption of small prey.[138] Leopards can live successfully in tiger habitat when there is abundant food and vegetation cover.[136][139] Otherwise, they appear to be less common where tigers are numerous. The recovery of the tiger population inRajaji National Park during the 2000s led to a reduction in leopard population densities.[140]
Reproduction and life cycle
Leopards mating
Leopard cubs in tree
In some areas, leopards mate all year round. InManchuria andSiberia, 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.[61] The female'sestrous cycle lasts about 46 days, and she is usually in heat for 6–7 days.[141]Gestation lasts for 90 to 105 days.[142] Cubs are usually born in alitter of 2–4 cubs.[143] The mortality rate of cubs is estimated at 41–50% during the first year.[112] Predators are the biggest cause for leopard cub mortality during their first year. Male leopards are known to causeinfanticide, in order to bring the female back into heat.[75] Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.[61]
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.[110][67] 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.[75] 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.[144] After separating from their mother, sibling cubs may travel together for months.[61] Both male and female leopards typically reach sexual maturity at 2–2⅓ years.[75]
Thegeneration length of the leopard is 9.3 years.[145]The average life span of a leopard is 12–17 years.[110] The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.[146]
Conservation
The leopard is listed onCITES 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 inSochi National Park; six individuals released intoCaucasus Nature Reserve andAlaniya National Park in 2018 survived as of February 2022.[147]
Threats
The leopard is primarily threatened byhabitat 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 bytrophy hunting andpoaching.[2] Contemporary records suggest that the leopard occurs in only 25% of its historical range.[81][148]
Between 2002 and 2012, at least four leopards were estimated to have been poached per week in India for theillegal wildlife trade of its skins and bones.[149]In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.[150] In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.[151]
Surveys in the Central African Republic'sChinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period,transhumantpastoralists from the border area with Sudan moved in the area with their livestock. Rangers confiscated large amounts ofpoison in the camps of livestock herders who were accompanied by armed merchants. They engaged in poaching large herbivores, sale ofbushmeat and trading leopard skins inAm Dafok.[152]
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.[153]
Leopards have been featured in art, mythology and folklore of many countries. InGreek mythology, it was a symbol of the godDionysus, who was depicted wearing leopard skin and using leopards as means of transportation. In one myth, the god was captured by pirates but two leopards rescued him.[154] Numerous Roman mosaics from North African sites depict fauna now found only intropical Africa.[155] During theBenin Empire, the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king oroba, since the leopard was considered the king of the forest.[156] TheAshanti 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.[154]
InRudyard Kipling's "How the Leopard Got His Spots", one of hisJust So Stories, a leopard with no spots in theHighveld 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.[157] A leopard played an important role in the 1938 Hollywood filmBringing Up Baby. African chiefs, European queens, Hollywood actors andburlesque dancers wore coats made of leopard skins.[154]
TheLeopard of Rudraprayag killed more than 125 people; the Panar Leopard was thought to have killed over 400 people. Both were shot by British hunterJim Corbett.[160] Thespotted devil of Gummalapur killed about 42 people in Karnataka, India.[161]
In captivity
Animal trainer with leopard
Theancient Romans kept leopards in captivity to be slaughtered inhunts as well asexecute criminals.[154] In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities.[156] Several leopards were kept in a menagerie originally established by KingJohn of England at theTower of London in the 13th century; around 1235, three of these animals were given toHenry III by Holy Roman EmperorFrederick II.[162] In modern times, leopards have beentrained andtamed incircuses.[154]
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