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Leopard

May 18, 2023

For other uses, see Leopard (disambiguation) and Leopards (disambiguation).

The leopard (Panthera pardus) is one of the five extant species in the genus Panthera. It occurs across a wide range in across Africa and Asia. It can adapt to a variety of habitats ranging from rainforest to steppe, including arid and montane areas. The leopard has relatively short legs and an elongated body with a large skull. Its fur is marked with rosettes. It is similar in appearance to the jaguar (Panthera onca) and both species are sometimes born melanistic, known as black panthers.

Leopard
Temporal range: Early Pleistocene – Present[2]
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Male African leopard in
Maasai Mara National Reserve, Kenya
CITES Appendix I (CITES)[3]
Scientific classification
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[3]

The leopard is an opportunistic predator, hunting a variety of species including ungulates and primates. It relies on its spotted pattern for camouflage as it stalks and ambushes its prey, which may be dragged up a tree. Throughout its range, it must deal with competition for other predators like lions and tigers. Leopards are solitary animals outside of mating and raising cubs.

It is listed as Vulnerable on the IUCN Red List because leopard populations are 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 have been 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.[4][5][6] The word λέοπάρδος originally referred to a cheetah (Acinonyx jubatus).[7]

'Panther' is another common name, derived from Latin: panther and Ancient Greek: πάνθηρ (pánthēr);[4] The generic name Panthera originates in Latin: panthera, a hunting net for catching wild beasts to be used by the Romans in combats.[8] Pardus is the masculine singular form.[9]

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.[10] 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.[11] 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.[12] 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 rain forest habitats.[13] Rosette patterns are unique in each individual.[14][15] This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as camouflage.[16]

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.[17] 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.[14][18] Its fur tends to grow longer in colder climates.[19] The leopard's rosettes differ from those of the jaguar (Panthera onca), which are darker and with smaller spots inside.[11] The leopard has a diploid chromosome number of 38.[20]

Melanistic leopards are also known as black panthers. Melanism in leopards is caused by a recessive allele and is inherited as a recessive trait.[21][22][23][24] In India, nine pale and white leopards were reported between 1905 and 1967.[25] 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.[26]

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.[17] 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).[27] 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.[18][19][28] The maximum recorded weight of a wild leopard in Southern Africa was around 96 kg (212 lb), and it measured 262 cm (103 in).[29] 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.[30][31]

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).[32]

Taxonomy

 
Map showing approximate distribution of leopard subspecies

Felis pardus was the scientific name proposed by Carl Linnaeus in 1758.[33] The generic name Panthera was first used by Lorenz Oken in 1816, who included all the known spotted cats into this group.[34] Oken's classification was not widely accepted, and Felis or Leopardus was used as the generic name until the early 20th century.[35]

The leopard was designated as the type species of Panthera by Joel Asaph Allen in 1902.[36] In 1917, Reginald Innes Pocock also subordinated the tiger (P. tigris), lion (P. leo), and jaguar (P. onca) to Panthera.[37][38]

Living subspecies

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.[39] Later analysis revealed a ninth valid subspecies, the Arabian leopard.[40]

In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:[41]

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.[3]  
Indian leopard (P. p. fusca) (Meyer, 1794)[42] It occurs in the Indian subcontinent, Myanmar and southern Tibet.[3][41][43]  
Javan leopard (P. p. melas) (Cuvier, 1809)[44] It is native to Java in Indonesia and is considered Critically Endangered.[3]  
Arabian leopard (P. p. nimr) (Hemprich and Ehrenberg, 1830)[45] It is native to the Arabian Peninsula, but considered locally extinct in the Sinai Peninsula. It is the smallest leopard subspecies.[46]  
Persian leopard (P. p. tulliana) (Valenciennes, 1856)[47] It is native to eastern Turkey, the Caucasus, southern Russia, the Iranian Plateau and the Hindu Kush. It is considered Endangered.[3]

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.[48]

  
Amur leopard (P. p. orientalis) (Schlegel, 1857)[49][50] It is native to the Russian Far East and northern China, but is locally extinct in the Korean peninsula.[3]  
Indochinese leopard (P. p. delacouri) Pocock, 1930[51] It occurs in mainland Southeast Asia and southern China.[3]  
Sri Lankan leopard (P. p. kotiya) Deraniyagala, 1956[52] It is native to Sri Lanka.[3]  

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.[53]

Evolution

 
Two cladograms proposed for Panthera. The upper cladogram is based on the 2006[54] and 2009[55] studies, while the lower is based on the 2010[56] and 2011[57] 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.[54][55]

Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion.[58] 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.[59] The leopard-lion clade diverged 3.1–1.95 million years ago.[56][57] 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.[60]

Fossils of leopard ancestors were excavated in East Africa and South Asia, dating back to the Pleistocene between 2 and 3.5 million years ago. The modern leopard is suggested to have evolved in Africa about 0.5 to 0.8 million years ago and to have radiated across Asia about 0.2 and 0.3 million years ago.[40] Fossil cat teeth collected in Sumatra's Padang Highlands were assigned to the leopard. It has since been hypothesized that it became extirpated on the island due to the Toba eruption about 75,000 years ago,[61] and due to competition with the Sunda clouded leopard (Neofelis diardi) and the dhole (Cuon alpinus).[62]

In Europe, the leopard occurred at least since the Pleistocene. 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,[63] Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central Poland.[64] The oldest known leopard fossils excavated in Europe are about 600,000 years old and were found in the Grotte du Vallonnet in France and near Mauer in Germany.[2] 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. The most recent, P. p. spelaea, appeared at the beginning of the Late Pleistocene and survived until about 24,000 years ago in several parts of Europe.[65] Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago.[66] Leopard fossils have also been found in Taiwan.[67]

Hybrids

Main articles: Panthera hybrid and Pumapard

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 were unsuccessful.[68]

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 extirpated in North Africa.[3] It inhabits foremost savanna and rainforest, and areas where grasslands, woodlands, and riverine forests remain largely undisturbed.[13] 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.[69]

Leopard populations in the Arabian Peninsula are small and fragmented.[70][71][72] In southeastern Egypt, a leopard killed in 2017 was the first sighting of the species in this area in 65 years.[73] In western and central Asia, it avoids deserts, areas with long snow cover and close proximity to urban centres.[74]

In the Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other Panthera species.[3] 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.[75] 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.[76]

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.[77] In Sri Lanka, leopards were recorded in Yala National Park and in unprotected forest patches, tea estates, grasslands, home gardens, pine and eucalyptus plantations.[78][79] In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's Karen State.[80] 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.[81] In Laos, leopards were recorded in Nam Et-Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area.[82][83] In Cambodia, leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest.[84][85] In southern China, leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009.[86]

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.[87]

In the Russian Far East, it inhabits temperate coniferous forests where winter temperatures reach a low of −25 °C (−13 °F).[40]

Behaviour and ecology

Leopard visual communication
 
A female leopard with white markings on the backs of her ears.
 
A female leopard showing white spots on the tail

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.[18] The roaring sequence in leopards consists mainly of grunts,[88] also called "sawing", as it resembles the sound of sawing wood. Cubs call their mother with an urr-urr sound.[18]

The whitish spots on the back of its ears are thought to play a role in communication.[89] 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.[90][91]

 
A leopard climbing down a tree

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.[18][28] In some regions, they are nocturnal.[92][93] 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.[94]

Video of a leopard in the wild

Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.[13] 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.[88]

Social spacing

In Kruger National Park, most leopards tend to keep 1 km (0.62 mi) apart.[95] Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.[96][97] Aggressive encounters are rare, typically limited to defending territories from intruders.[19] In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.[92]

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.[98] 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.[18][28]

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.[99] Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the Serengeti, males have home ranges of 33–38 km2 (13–15 sq mi) and females of 14–16 km2 (5.4–6.2 sq mi);[100][101] but males in northeastern Namibia of 451 km2 (174 sq mi) and females of 188 km2 (73 sq mi).[102] They are even larger in arid and montane areas.[19] In Nepal's 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.[103]

Hunting and diet

Stages of the hunt
 
Stalking
 
Killing a young bushbuck
 
Dragging an impala kill
 
Caching the kill up a tree

The leopard is a carnivore 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 are ungulates, such as impala (Aepyceros melampus), bushbuck (Tragelaphus scriptus), common duiker (Sylvicapra grimmia) and chital (Axis axis). Primates preyed upon include white-eyelid mangabeys (Cercocebus sp.), guenons (Cercopithecus sp.) and gray langurs (Semnopithecus sp.). Leopards also kill smaller carnivores like black-backed jackal (Lupulella mesomelas), bat-eared fox (Otocyon megalotis), genet (Genetta sp.) and cheetah.[104]

The largest prey killed by a leopard was reportedly a male eland weighing 900 kg (2,000 lb).[88] 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 (Elaphodus cephalophus) to pursuing bamboo rats (Rhizomys sinense) and other smaller prey.[105]

The leopard depends mainly on its acute senses of hearing and vision for hunting.[106] It primarily hunts at night in most areas.[18] In western African forests and Tsavo National Park, they have also been observed hunting by day.[107] They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.[108]

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.[18] It caches kills up to 2 km (1.2 mi) apart.[96] 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.[107] 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.[19][109]

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.[95] In the southern Kalahari Desert, leopards meet their water requirements by the bodily fluids of prey and succulent plants; they drink water every two to three days and feed infrequently on moisture-rich plants such as gemsbok cucumbers (Acanthosicyos naudinianus), watermelon (Citrullus lanatus) and Kalahari sour grass (Schmidtia kalahariensis).[110]

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 (Panthera tigris), lion (P. leo), cheetah (Acinonyx jubatus), spotted hyena (Crocuta crocuta), striped hyena (Hyaena hyaena), brown hyena (Parahyaena brunnea), African wild dog (Lycaon pictus), dhole (Cuon alpinus), wolf (Canis lupus) 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 (Civettictis civetta), caracal (Caracal caracal) and African wildcat (Felis lybica).[13][111] 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 (Melursus ursinus) apparently result in both animals winding up either dead or grievously injured.[112][113] Leopards generally avoid large packs of African wild dogs and dholes and will flee up a tree at the sight of them.[114][115]

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.[108][111][116][117] In the Kalahari Desert, leopards frequently lose kills to brown hyenas, if their unable to move the kill up a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills.[118][119] Lions occasionally fetch leopard kills from trees.[109]

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.[13] In areas where leopards and tigers are sympatric, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous.[116] Tigers appear to inhabit the deep parts of the forest while leopards are pushed closer to the fringes.[120] 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 that may be more prevalent in the leopard's co-existence with the lion in savanna habitats.[121]

Nile crocodiles (Crocodylus niloticus) 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.[95][96] Mugger crocodiles (C. palustris) reportedly killed an adult leopard in Rajasthan.[122] An adult leopard was recovered from the stomach of a 5.5 m (18 ft 1 in) Burmese python (Python bivittatus).[123] In the Serengeti National Park, troops of around 30–40 olive baboons (Papio anubis) were observed mobbing and attacking a female leopard and her cubs.[124]

Reproduction and life cycle

 
A female in estrus fights with a male attempting to mate with her
 
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.[125] The female's estrous cycle lasts about 46 days, and she is usually in heat for 6–7 days.[126] The generation length of the leopard is 9.3 years.[127] Gestation lasts for 90 to 105 days.[128] Cubs are usually born in a litter of 2–4 cubs.[129] The mortality rate of cubs is estimated at 41–50% during the first year.[95] 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.[130] Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.[125]

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.[88][19] 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.[130] 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.[131] After separating from their mother, sibling cubs may travel together for months.[125] Both male and female leopards typically reach sexual maturity at 2–2⅓ years.[130]

The average life span of a leopard is 12–17 years.[88] The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.[132]

Conservation issues

The leopard is listed on CITES Appendix I, and trade is restricted to skins and body parts of 2,560 individuals in 11 sub-Saharan countries.[3] 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.[3] Contemporary records suggest that the leopard occurs in only 25% of its historical range.[133][134]

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.[135] In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.[136] In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.[137]

Surveys in the Central African Republic's Chinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period, 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.[138]

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.[139]

The leopard is considered locally extinct in Hong Kong, Singapore, South Korea, Jordan, Morocco, Togo, the United Arab Emirates, Uzbekistan, Lebanon, Mauritania, Kuwait, Syria, Libya, Tunisia and most likely in North Korea, Gambia, Laos, Lesotho, Tajikistan, Vietnam and Israel.[3]

Human interaction

Cultural significance

 
Leopards on the Magerius Mosaic from modern Tunisia. Numerous Roman mosaics from North African sites depict fauna now found only in tropical Africa.[140]
 
Rock art of P. pardus spelaea in Chauvet cave
 
Leopard head to hip ornament from the Court of Benin

Leopards have been featured in art, mythology and folklore of many countries. In Greek mythology, it was a symbol of the god Dionysus, 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.[141] During the Benin Empire, the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king or oba, since the leopard was considered the king of the forest.[142] The Ashanti 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.[141]

In Rudyard Kipling's "How the Leopard Got His Spots", one of his Just So Stories, a leopard with no spots in the High Veldt 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.[143] 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.[141]

The leopard is a frequently used in heraldry, most commonly as passant.[144] The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the heraldic lion, and the two are often used interchangeably. Naturalistic leopard-like depictions appear on the coat of arms of Benin, Malawi, Somalia, the Democratic Republic of the Congo and Gabon, the last of which uses a black panther.[145]

Attacks on people

Main article: Leopard attack

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.[146] The spotted devil of Gummalapur killed about 42 people in Karnataka, India.[147]

In captivity

 
Animal trainer with leopard

The Ancient Romans kept leopards in captivity to be slaughtered in hunts as well as to be used in executions of criminals.[141] In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities.[142] Several leopards were kept in a menagerie established by King 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.[148] In modern times, leopards have been trained and tamed in circuses.[141]

See also

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Further reading

  • Allsen, Thomas T. (2007). "Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries". In Mair, Victor H. (ed.). Contact and Exchange in the Ancient World. Honolulu: University of Hawai'i Press. ISBN 978-0-8248-2884-4.
  • DeRuiter, D. J.; Berger, L. R. (2000). "Leopards as Taphonomic Agents in dolomitic Caves—Implications for bone Accumulations in the Hominid-bearing Deposits of South Africa". Journal of Archaeological Science. 27 (8): 665–684. doi:10.1006/jasc.1999.0470.
  • Schaller, G. B. (1972). The Serengeti Lion. Chicago: University of Chicago Press. ISBN 978-0-226-73639-6.
  • Sanei, A. (2007). Analysis of leopard (Panthera pardus) status in Iran (in Persian). Tehran: Sepehr Publication Center. ISBN 978-964-6123-74-8.
  • Sanei, A.; Zakaria, M.; Yusof, E.; Roslan, M. (2011). "Estimation of leopard population size in a secondary forest within Malaysia's capital agglomeration using unsupervised classification of pugmarks" (PDF). Tropical Ecology. 52 (1): 209–217. (PDF) from the original on 2011-10-02.
  • Taylor, P.; Barrientos, S.; Dolan, C. (2005). Beyond Conservation: A Wildland Strategy. Earthscan. ISBN 978-1-84407-197-5.
  • Zakaria, M.; Sanei, A. (2011). "Conservation and management prospects of the Persian and Malayan leopards". Asia Life Sciences. Supplement 7: 1–5.

External links

 
Wikimedia Commons has media related to:
Panthera pardus (category)
  • IUCN/SSC Cat Specialist Group: Panthera pardus in Africa and Panthera pardus in Asia
  • "Leopard" . Encyclopædia Britannica (11th ed.). 1911.

May 18, 2023
leopard, other, uses, disambiguation, disambiguation, leopard, panthera, pardus, five, extant, species, genus, panthera, occurs, across, wide, range, across, africa, asia, adapt, variety, habitats, ranging, from, rainforest, steppe, including, arid, montane, a. For other uses see Leopard disambiguation and Leopards disambiguation The leopard Panthera pardus is one of the five extant species in the genus Panthera It occurs across a wide range in across Africa and Asia It can adapt to a variety of habitats ranging from rainforest to steppe including arid and montane areas The leopard has relatively short legs and an elongated body with a large skull Its fur is marked with rosettes It is similar in appearance to the jaguar Panthera onca and both species are sometimes born melanistic known as black panthers LeopardTemporal range Early Pleistocene Present 2 PreꞒ Ꞓ O S D C P T J K Pg N Male African leopard in Maasai Mara National Reserve KenyaConservation statusVulnerable IUCN 3 1 3 CITES Appendix I CITES 3 Scientific classificationKingdom AnimaliaPhylum ChordataClass MammaliaOrder CarnivoraSuborder FeliformiaFamily FelidaeSubfamily PantherinaeGenus PantheraSpecies P pardus 1 Binomial namePanthera pardus 1 Linnaeus 1758 SubspeciesSee textPresent and historical distribution of the leopard 3 The leopard is an opportunistic predator hunting a variety of species including ungulates and primates It relies on its spotted pattern for camouflage as it stalks and ambushes its prey which may be dragged up a tree Throughout its range it must deal with competition for other predators like lions and tigers Leopards are solitary animals outside of mating and raising cubs It is listed as Vulnerable on the IUCN Red List because leopard populations are 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 have been popular in fashion Contents 1 Etymology 2 Characteristics 2 1 Size 3 Taxonomy 3 1 Living subspecies 3 2 Evolution 3 3 Hybrids 4 Distribution and habitat 5 Behaviour and ecology 5 1 Social spacing 5 2 Hunting and diet 5 3 Enemies and competitors 5 4 Reproduction and life cycle 6 Conservation issues 7 Human interaction 7 1 Cultural significance 7 2 Attacks on people 7 3 In captivity 8 See also 9 References 10 Further reading 11 External linksEtymologyThe English name leopard comes from Old French leupart or Middle French liepart that derives from Latin leopardus and Ancient Greek leopardos leopardos Leopardos could be a compound of lewn leōn meaning lion and pardos pardos meaning spotted 4 5 6 The word leopardos originally referred to a cheetah Acinonyx jubatus 7 Panther is another common name derived from Latin panther and Ancient Greek pan8hr panther 4 The generic name Panthera originates in Latin panthera a hunting net for catching wild beasts to be used by the Romans in combats 8 Pardus is the masculine singular form 9 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 10 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 11 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 12 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 rain forest habitats 13 Rosette patterns are unique in each individual 14 15 This pattern is thought to be an adaptation to dense vegetation with patchy shadows where it serves as camouflage 16 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 17 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 14 18 Its fur tends to grow longer in colder climates 19 The leopard s rosettes differ from those of the jaguar Panthera onca which are darker and with smaller spots inside 11 The leopard has a diploid chromosome number of 38 20 Melanistic leopards are also known as black panthers Melanism in leopards is caused by a recessive allele and is inherited as a recessive trait 21 22 23 24 In India nine pale and white leopards were reported between 1905 and 1967 25 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 26 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 17 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 27 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 18 19 28 The maximum recorded weight of a wild leopard in Southern Africa was around 96 kg 212 lb and it measured 262 cm 103 in 29 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 30 31 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 32 Taxonomy Map showing approximate distribution of leopard subspecies Felis pardus was the scientific name proposed by Carl Linnaeus in 1758 33 The generic name Panthera was first used by Lorenz Oken in 1816 who included all the known spotted cats into this group 34 Oken s classification was not widely accepted and Felis or Leopardus was used as the generic name until the early 20th century 35 The leopard was designated as the type species of Panthera by Joel Asaph Allen in 1902 36 In 1917 Reginald Innes Pocock also subordinated the tiger P tigris lion P leo and jaguar P onca to Panthera 37 38 Living subspecies 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 39 Later analysis revealed a ninth valid subspecies the Arabian leopard 40 In 2017 the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa 41 Subspecies Distribution ImageAfrican leopard P p pardus Linnaeus 1758 1 It is the most widespread leopard subspecies and is native to most of Sub Saharan Africa 3 Indian leopard P p fusca Meyer 1794 42 It occurs in the Indian subcontinent Myanmar and southern Tibet 3 41 43 Javan leopard P p melas Cuvier 1809 44 It is native to Java in Indonesia and is considered Critically Endangered 3 Arabian leopard P p nimr Hemprich and Ehrenberg 1830 45 It is native to the Arabian Peninsula but considered locally extinct in the Sinai Peninsula It is the smallest leopard subspecies 46 Persian leopard P p tulliana Valenciennes 1856 47 It is native to eastern Turkey the Caucasus southern Russia the Iranian Plateau and the Hindu Kush It is considered Endangered 3 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 48 Amur leopard P p orientalis Schlegel 1857 49 50 It is native to the Russian Far East and northern China but is locally extinct in the Korean peninsula 3 Indochinese leopard P p delacouri Pocock 1930 51 It occurs in mainland Southeast Asia and southern China 3 Sri Lankan leopard P p kotiya Deraniyagala 1956 52 It is native to Sri Lanka 3 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 53 Evolution Two cladograms proposed for Panthera The upper cladogram is based on the 2006 54 and 2009 55 studies while the lower is based on the 2010 56 and 2011 57 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 54 55 Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion 58 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 59 The leopard lion clade diverged 3 1 1 95 million years ago 56 57 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 60 Fossils of leopard ancestors were excavated in East Africa and South Asia dating back to the Pleistocene between 2 and 3 5 million years ago The modern leopard is suggested to have evolved in Africa about 0 5 to 0 8 million years ago and to have radiated across Asia about 0 2 and 0 3 million years ago 40 Fossil cat teeth collected in Sumatra s Padang Highlands were assigned to the leopard It has since been hypothesized that it became extirpated on the island due to the Toba eruption about 75 000 years ago 61 and due to competition with the Sunda clouded leopard Neofelis diardi and the dhole Cuon alpinus 62 In Europe the leopard occurred at least since the Pleistocene 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 Prerov in the Czech Republic and the Baranya in southern Hungary 63 Leopard fossils dating to the Late Pleistocene were found in Bisnik Cave in south central Poland 64 The oldest known leopard fossils excavated in Europe are about 600 000 years old and were found in the Grotte du Vallonnet in France and near Mauer in Germany 2 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 The most recent P p spelaea appeared at the beginning of the Late Pleistocene and survived until about 24 000 years ago in several parts of Europe 65 Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago 66 Leopard fossils have also been found in Taiwan 67 Hybrids Main articles Panthera hybrid and Pumapard 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 were unsuccessful 68 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 extirpated in North Africa 3 It inhabits foremost savanna and rainforest and areas where grasslands woodlands and riverine forests remain largely undisturbed 13 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 69 Leopard populations in the Arabian Peninsula are small and fragmented 70 71 72 In southeastern Egypt a leopard killed in 2017 was the first sighting of the species in this area in 65 years 73 In western and central Asia it avoids deserts areas with long snow cover and close proximity to urban centres 74 In the Indian subcontinent the leopard is still relatively abundant with greater numbers than those of other Panthera species 3 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 75 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 76 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 77 In Sri Lanka leopards were recorded in Yala National Park and in unprotected forest patches tea estates grasslands home gardens pine and eucalyptus plantations 78 79 In Myanmar leopards were recorded for the first time by camera traps in the hill forests of Myanmar s Karen State 80 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 81 In Laos leopards were recorded in Nam Et Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area 82 83 In Cambodia leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest 84 85 In southern China leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009 86 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 87 In the Russian Far East it inhabits temperate coniferous forests where winter temperatures reach a low of 25 C 13 F 40 Behaviour and ecologyLeopard visual communication A female leopard with white markings on the backs of her ears A female leopard showing white spots on the tail 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 18 The roaring sequence in leopards consists mainly of grunts 88 also called sawing as it resembles the sound of sawing wood Cubs call their mother with an urr urr sound 18 The whitish spots on the back of its ears are thought to play a role in communication 89 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 90 91 A leopard climbing down a tree 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 18 28 In some regions they are nocturnal 92 93 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 94 source source source source source source Video of a leopard in the wild Leopards can climb trees quite skillfully often resting on tree branches and descending headfirst 13 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 88 Social spacing In Kruger National Park most leopards tend to keep 1 km 0 62 mi apart 95 Males occasionally interact with their partners and cubs and exceptionally this can extend beyond to two generations 96 97 Aggressive encounters are rare typically limited to defending territories from intruders 19 In a South African reserve a male was wounded in a male male territorial battle over a carcass 92 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 98 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 18 28 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 99 Sizes of home ranges vary geographically and depending on habitat and availability of prey In the Serengeti males have home ranges of 33 38 km2 13 15 sq mi and females of 14 16 km2 5 4 6 2 sq mi 100 101 but males in northeastern Namibia of 451 km2 174 sq mi and females of 188 km2 73 sq mi 102 They are even larger in arid and montane areas 19 In Nepal s 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 103 Hunting and diet Stages of the hunt Stalking Killing a young bushbuck Dragging an impala kill Caching the kill up a tree The leopard is a carnivore 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 are ungulates such as impala Aepyceros melampus bushbuck Tragelaphus scriptus common duiker Sylvicapra grimmia and chital Axis axis Primates preyed upon include white eyelid mangabeys Cercocebus sp guenons Cercopithecus sp and gray langurs Semnopithecus sp Leopards also kill smaller carnivores like black backed jackal Lupulella mesomelas bat eared fox Otocyon megalotis genet Genetta sp and cheetah 104 The largest prey killed by a leopard was reportedly a male eland weighing 900 kg 2 000 lb 88 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 Elaphodus cephalophus to pursuing bamboo rats Rhizomys sinense and other smaller prey 105 The leopard depends mainly on its acute senses of hearing and vision for hunting 106 It primarily hunts at night in most areas 18 In western African forests and Tsavo National Park they have also been observed hunting by day 107 They usually hunt on the ground In the Serengeti they have been seen to ambush prey by descending on it from trees 108 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 18 It caches kills up to 2 km 1 2 mi apart 96 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 107 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 19 109 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 95 In the southern Kalahari Desert leopards meet their water requirements by the bodily fluids of prey and succulent plants they drink water every two to three days and feed infrequently on moisture rich plants such as gemsbok cucumbers Acanthosicyos naudinianus watermelon Citrullus lanatus and Kalahari sour grass Schmidtia kalahariensis 110 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 Panthera tigris lion P leo cheetah Acinonyx jubatus spotted hyena Crocuta crocuta striped hyena Hyaena hyaena brown hyena Parahyaena brunnea African wild dog Lycaon pictus dhole Cuon alpinus wolf Canis lupus 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 Civettictis civetta caracal Caracal caracal and African wildcat Felis lybica 13 111 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 Melursus ursinus apparently result in both animals winding up either dead or grievously injured 112 113 Leopards generally avoid large packs of African wild dogs and dholes and will flee up a tree at the sight of them 114 115 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 108 111 116 117 In the Kalahari Desert leopards frequently lose kills to brown hyenas if their unable to move the kill up a tree Single brown hyenas have been observed charging at and displacing male leopards from kills 118 119 Lions occasionally fetch leopard kills from trees 109 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 13 In areas where leopards and tigers are sympatric coexistence is reportedly not the general rule with leopards being few where tigers are numerous 116 Tigers appear to inhabit the deep parts of the forest while leopards are pushed closer to the fringes 120 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 that may be more prevalent in the leopard s co existence with the lion in savanna habitats 121 Nile crocodiles Crocodylus niloticus 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 95 96 Mugger crocodiles C palustris reportedly killed an adult leopard in Rajasthan 122 An adult leopard was recovered from the stomach of a 5 5 m 18 ft 1 in Burmese python Python bivittatus 123 In the Serengeti National Park troops of around 30 40 olive baboons Papio anubis were observed mobbing and attacking a female leopard and her cubs 124 Reproduction and life cycle A female in estrus fights with a male attempting to mate with her 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 2 and three and males between the ages of two and three 125 The female s estrous cycle lasts about 46 days and she is usually in heat for 6 7 days 126 The generation length of the leopard is 9 3 years 127 Gestation lasts for 90 to 105 days 128 Cubs are usually born in a litter of 2 4 cubs 129 The mortality rate of cubs is estimated at 41 50 during the first year 95 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 130 Intervals between births average 15 to 24 months but can be shorter depending on the survival of the cubs 125 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 88 19 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 130 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 131 After separating from their mother sibling cubs may travel together for months 125 Both male and female leopards typically reach sexual maturity at 2 2 years 130 The average life span of a leopard is 12 17 years 88 The oldest leopard was a captive female that died at the age of 24 years 2 months and 13 days 132 Conservation issuesThe leopard is listed on CITES Appendix I and trade is restricted to skins and body parts of 2 560 individuals in 11 sub Saharan countries 3 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 3 Contemporary records suggest that the leopard occurs in only 25 of its historical range 133 134 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 135 In spring 2013 37 leopard skins were found during a 7 week long market survey in major Moroccan cities 136 In 2014 43 leopard skins were detected during two surveys in Morocco Vendors admitted to have imported skins from sub Saharan Africa 137 Surveys in the Central African Republic s Chinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017 In this period 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 138 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 139 The leopard is considered locally extinct in Hong Kong Singapore South Korea Jordan Morocco Togo the United Arab Emirates Uzbekistan Lebanon Mauritania Kuwait Syria Libya Tunisia and most likely in North Korea Gambia Laos Lesotho Tajikistan Vietnam and Israel 3 Human interactionCultural significance Leopards on the Magerius Mosaic from modern Tunisia Numerous Roman mosaics from North African sites depict fauna now found only in tropical Africa 140 Rock art of P pardus spelaea in Chauvet cave Leopard head to hip ornament from the Court of Benin Leopards have been featured in art mythology and folklore of many countries In Greek mythology it was a symbol of the god Dionysus 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 141 During the Benin Empire the leopard was commonly represented on engravings and sculptures and was used to symbolise the power of the king or oba since the leopard was considered the king of the forest 142 The Ashanti 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 141 In Rudyard Kipling s How the Leopard Got His Spots one of his Just So Stories a leopard with no spots in the High Veldt 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 143 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 141 The leopard is a frequently used in heraldry most commonly as passant 144 The heraldic leopard lacks spots and sports a mane making it visually almost identical to the heraldic lion and the two are often used interchangeably Naturalistic leopard like depictions appear on the coat of arms of Benin Malawi Somalia the Democratic Republic of the Congo and Gabon the last of which uses a black panther 145 Attacks on people Main article Leopard attack 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 146 The spotted devil of Gummalapur killed about 42 people in Karnataka India 147 In captivity Animal trainer with leopard The Ancient Romans kept leopards in captivity to be slaughtered in hunts as well as to be used in executions of criminals 141 In Benin leopards were kept and paraded as mascots totems and sacrifices to deities 142 Several leopards were kept in a menagerie established by King 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 148 In modern times leopards have been trained and tamed in circuses 141 See alsoLeopard pattern List of largest cats Panther legendary creature References a b Wozencraft W C 2005 Species Panthera pardus In Wilson D E Reeder D M eds Mammal Species of the World A Taxonomic and Geographic Reference 3rd ed Johns Hopkins University Press p 547 ISBN 978 0 8018 8221 0 OCLC 62265494 a b Ghezzo E amp Rook L 2015 The remarkable Panthera pardus Felidae Mammalia record from Equi Massa Italy taphonomy morphology and paleoecology Quaternary Science Reviews 110 110 131 151 doi 10 1016 j quascirev 2014 12 020 a b c d e f g h i j k l m n o Stein A B Athreya V Gerngross P Balme G Henschel P Karanth U Miquelle D Rostro Garcia S Kamler J F Laguardia A Khorozyan I amp Ghoddousi A 2020 amended version of 2019 assessment Panthera pardus IUCN Red List of Threatened Species 2020 e T15954A163991139 doi 10 2305 IUCN UK 2020 1 RLTS T15954A163991139 en Retrieved 15 January 2022 a b Lewis C T amp Short C 1879 leǒpardus A Latin Dictionary Oxford Clarendon Press p 1069 Liddell H G amp Scott R 1889 leo pardos A Greek English Lexicon Oxford Clarendon Press p 884 Partridge E 1983 Origins A Short Etymological Dictionary of Modern English New York Greenwich House p 349 ISBN 978 0 517 41425 5 Nicholas N 1999 A conundrum of cats pards and their relatives in Byzantium Greek Roman and Byzantine Studies 40 253 298 S2CID 56160515 Lewis C T amp Short C 1879 panthera A Latin Dictionary Oxford Clarendon Press p 1298 Lewis C T amp Short C 1879 pardus A Latin Dictionary Oxford Clarendon Press p 1302 Mills M G L 2005 Subfamily Pantherinae In Skinner J D Chimimba C T eds The mammals of the southern African sub region Third ed Cambridge Cambridge University Press pp 385 396 ISBN 9780521844185 a b Mivart St G J 1900 Different kind of Cats The Cat An Introduction to the Study of Backboned Animals Especially Mammals London John Murray pp 391 439 Pocook R I 1932 The Leopards of Africa Proceedings of the Zoological Society of London 102 2 543 591 doi 10 1111 j 1096 3642 1932 tb01085 x a b c d e Nowell K amp Jackson P 1996 Leopard Panthera pardus Linnaeus 1758 Wild Cats status survey and conservation action plan Gland Switzerland IUCN SSC Cat Specialist Group Archived from the original on 2014 02 22 a b Schutze H 2002 Field Guide to the Mammals of the Kruger National Park Cape Town South Africa Struik Publishers pp 92 93 ISBN 978 1 86872 594 6 Menon V 2014 Indian Mammals A Field Guide Gurgaon India Hachette ISBN 978 93 5009 761 8 Allen W L Cuthill I C Scott Samuel N E amp Baddeley R 2010 Why the leopard got its spots relating pattern development to ecology in felids Proceedings of the Royal Society B 278 1710 1373 1380 doi 10 1098 rspb 2010 1734 PMC 3061134 PMID 20961899 a b Hoath R 2009 Leopard Panthera pardus Linnaeus 1758 Field Guide to the Mammals of Egypt Cairo Egypt American University in Cairo Press pp 106 107 ISBN 978 977 416 254 1 a b c d e f g h Estes R 1991 Leopard Panthera pardus The Behavior Guide to African Mammals Including Hoofed Mammals Carnivores Primates Los Angeles The University of California Press pp 366 369 ISBN 978 0 520 08085 0 a b c d e f Stein A B amp Hayssen V 2010 Panthera pardus Carnivora Felidae Mammalian Species 45 900 30 48 doi 10 1644 900 1 S2CID 44839740 Heptner V G amp Sludskii A A 1992 1972 Bars leopard Mlekopitajuscie Sovetskogo Soiuza Moskva Vyssaia Skola Mammals of the Soviet Union Volume II Part 2 Washington DC Smithsonian Institution and the National Science Foundation pp 203 273 ISBN 978 90 04 08876 4 Robinson R 1970 Inheritance of the black form of the leopard Panthera pardus Genetica 41 1 190 197 doi 10 1007 BF00958904 PMID 5480762 S2CID 5446868 Eizirik E Yuhki N Johnson W E Menotti Raymond M Hannah S S O Brien S J 2003 Molecular genetics and evolution of melanism in the cat family Current Biology 13 5 448 453 doi 10 1016 S0960 9822 03 00128 3 PMID 12620197 S2CID 19021807 Kawanishi K Sunquist M E Eizirik E Lynam A J Ngoprasert D Wan Shahruddin W N Rayan D M Sharma D S K amp Steinmetz R 2010 Near fixation of melanism in leopards of the Malay Peninsula Journal of Zoology 282 3 201 206 doi 10 1111 j 1469 7998 2010 00731 x da Silva L G K Kawanishi K Henschel P Kittle A Sanei A Reebin A Miquelle D Stein A B Watson A Kekule L B Machado R B amp Eizirik E 2017 Mapping black panthers Macroecological modeling of melanism in leopards Panthera pardus PLOS One 12 4 e0170378 Bibcode 2017PLoSO 1270378D doi 10 1371 journal pone 0170378 PMC 5381760 PMID 28379961 Divyabhanusinh 1993 On mutant leopards Panthera pardus from India Journal of the Bombay Natural History Society 90 1 88 89 Pirie T J Thomas R L amp Fellowes M D E 2016 Erythristic leopards Panthera pardus in South Africa Bothalia 46 1 1 5 doi 10 4102 abc v46i1 2034 Kingdon Jonathan Happold David Butynski Thomas Hoffmann Michael Happold Meredith Kalina Jan 2013 05 23 Mammals of Africa A amp C Black p 168 ISBN 978 1 4081 8996 2 a b c Nowak R M 1999 Panthera pardus Leopard Walker s Mammals of the World Sixth ed Baltimore US Johns Hopkins University Press pp 828 831 ISBN 978 0 8018 5789 8 Burnie D amp Wilson D E eds 2001 Animal The Definitive Visual Guide to the World s Wildlife DK Adult ISBN 978 0 7894 7764 4 Is this the longest leopard in India The Times of India 2016 Leopard shot in Bilaspur turns out to be a record breaker The Tribune Trust 2016 Prater S H 1921 Record Panther Skull P p pardus The Journal of the Bombay Natural History Society XXVII Part IV 933 935 Linnaeus C 1758 Felis pardus Caroli Linnaei Systema naturae per regna tria naturae secundum classes ordines genera species cum characteribus differentiis synonymis locis Vol Tomus I decima reformata ed Holmiae Laurentius Salvius p 41 42 in Latin Oken L 1816 1 Art Panthera Lehrbuch der Zoologie 2 Abtheilung Jena August Schmid amp Comp p 1052 Ellerman J R Morrison Scott T C S 1966 Checklist of Palaearctic and Indian mammals 1758 to 1946 Second ed London British Museum of Natural History pp 315 317 Allen J A 1902 Mammal names proposed by Oken in his Lehrbuch der Zoologie PDF Bulletin of the American Museum of Natural History 16 27 373 379 Pocock R I 1917 The Classification of existing Felidae The Annals and Magazine of Natural History Series 8 XX 329 350 doi 10 1080 00222931709487018 Pocock R I 1939 Panthera pardus The Fauna of British India including Ceylon and Burma Mammalia Volume 1 London Taylor and Francis pp 222 239 Miththapala S Seidensticker J amp O Brien S J 1996 Phylogeographic subspecies recognition in leopards Panthera pardus molecular genetic variation PDF Conservation Biology 10 4 1115 1132 doi 10 1046 j 1523 1739 1996 10041115 x a b c Uphyrkina O Johnson E W Quigley H Miquelle D Marker L Bush M amp O Brien S J 2001 Phylogenetics genome diversity and origin of modern leopard Panthera pardus PDF Molecular Ecology 10 11 2617 2633 doi 10 1046 j 0962 1083 2001 01350 x PMID 11883877 S2CID 304770 Archived PDF from the original on 2011 09 10 a b Kitchener A C Breitenmoser Wursten C Eizirik E Gentry A Werdelin L Wilting A Yamaguchi N Abramov A V Christiansen P Driscoll C Duckworth J W Johnson W Luo S J Meijaard E O Donoghue P Sanderson J Seymour K Bruford M Groves C Hoffmann M Nowell K Timmons Z amp Tobe S 2017 A revised taxonomy of the Felidae The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group PDF Cat News Special Issue 11 73 75 Meyer F A A 1794 Uber de la Metheries schwarzen Panther Zoologische Annalen Erster Band Weimar Im Verlage des Industrie Comptoirs pp 394 396 Laguardia A Kamler J F Li S Zhang C Zhou Z amp Shi K 2017 The current distribution and status of leopards Panthera pardus in China Oryx 51 1 153 159 doi 10 1017 S0030605315000988 Cuvier G 1809 Recherches sur les especes vivantes de grands chats pour servir de preuves et d eclaircissement au chapitre sur les carnassiers fossils Annales du Museum National d Histoire Naturelle Tome XIV 136 164 Hemprich W Ehrenberg C G 1830 Felis pardus nimr In Dr C G Ehrenberg ed Symbolae Physicae seu Icones et Descriptiones Mammalium quae ex Itinere per Africam Borealem et Asiam Occidentalem Friderici Guilelmi Hemprich et Christiani Godofredi Ehrenberg Decas Secunda Zoologica I Mammalia II Berolini Officina Academica pp Plate 17 Spalton J A amp Al Hikmani H M 2006 The Leopard in the Arabian Peninsula Distribution and Subspecies Status PDF Cat News Special Issue 1 4 8 Archived PDF from the original on 2015 06 19 Valenciennes A 1856 Sur une nouvelles espece de Panthere tue par M Tchihatcheff a Ninfi village situe a huit lieues est de Smyrne Comptes Rendus Hebdomadaires des Seances de l Academie des Sciences 42 1035 1039 Khorozyan I G Gennady F Baryshnikov G F amp Abramov A V 2006 Taxonomic status of the leopard Panthera pardus Carnivora Felidae in the Caucasus and adjacent areas Russian Journal of Theriology 5 1 41 52 doi 10 15298 rusjtheriol 05 1 06 Schlegel H 1857 Felis orientalis Handleiding Tot de Beoefening der Dierkunde Ie Deel Breda Boekdrukkerij van Nys p 23 Gray J E 1862 Description of some new species of Mammalia Proceedings of the Royal Zoological Society of London 30 261 263 plate XXXIII doi 10 1111 j 1469 7998 1862 tb06524 x Pocock R I 1930 The Panthers and Ounces of Asia Journal of the Bombay Natural History Society 34 2 307 336 Deraniyagala P E P 1956 The Ceylon leopard a distinct subspecies Spolia Zeylanica 28 115 116 Anco C Kolokotronis S O Henschel P Cunningham S W Amato G amp Hekkala E 2017 Historical mitochondrial diversity in African leopards Panthera pardus revealed by archival museum specimens Mitochondrial DNA Part A 29 3 455 473 doi 10 1080 24701394 2017 1307973 PMID 28423965 S2CID 4348541 a b Johnson W E Eizirik E Pecon Slattery J Murphy W J Antunes A Teeling E amp O Brien S J 2006 The late Miocene radiation of modern Felidae a genetic assessment Science 311 5757 73 77 Bibcode 2006Sci 311 73J doi 10 1126 science 1122277 PMID 16400146 S2CID 41672825 a b Werdelin L Yamaguchi N Johnson W E amp O Brien S J 2010 Phylogeny and evolution of cats Felidae In Macdonald D W amp Loveridge A J eds Biology and Conservation of Wild Felids Oxford UK Oxford University Press pp 59 82 ISBN 978 0 19 923445 5 a b Davis B W Li G amp Murphy W J 2010 Supermatrix and species tree methods resolve phylogenetic relationships within the big cats Panthera Carnivora Felidae PDF Molecular Phylogenetics and Evolution 56 1 64 76 doi 10 1016 j ympev 2010 01 036 PMID 20138224 Archived from the original PDF on 2016 03 05 a b Mazak J H Christiansen P Kitchener A C amp Goswami A 2011 Oldest known pantherine skull and evolution of the tiger PLOS ONE 6 10 e25483 Bibcode 2011PLoSO 625483M doi 10 1371 journal pone 0025483 PMC 3189913 PMID 22016768 Bininda Emonds O R P Decker Flum D M amp Gittleman J L 2001 The utility of chemical signals as phylogenetic characters an example from the Felidae Biological Journal of the Linnean Society 72 1 1 15 doi 10 1111 j 1095 8312 2001 tb01297 x Tseng Z J Wang X Slater G J Takeuchi G T Li Q Liu J amp Xie G 2014 Himalayan fossils of the oldest known pantherine establish ancient origin of big cats Proceedings of the Royal Society B Biological Sciences 281 1774 20132686 doi 10 1098 rspb 2013 2686 PMC 3843846 PMID 24225466 Li G Davis B W Eizirik E amp Murphy W J 2016 Phylogenomic evidence for ancient hybridization in the genomes of living cats Felidae Genome Research 26 1 1 11 doi 10 1101 gr 186668 114 PMC 4691742 PMID 26518481 Wilting A Patel R Pfestorf H Kern C Sultan K Ario A Penaloza F Kramer Schadt S Radchuk V Foerster D W amp Fickel J 2016 Evolutionary history and conservation significance of the Javan leopard Panthera pardus melas Journal of Zoology 299 4 239 250 doi 10 1111 jzo 12348 Volmer R Holzchen E Wurster A Ferreras M R amp Hertler C 2017 Did Panthera pardus Linnaeus 1758 become extinct in Sumatra because of competition for prey Modeling interspecific competition within the Late Pleistocene carnivore guild of the Padang Highlands Sumatra Palaeogeography Palaeoclimatology Palaeoecology 487 175 186 Bibcode 2017PPP 487 175V doi 10 1016 j palaeo 2017 08 032 Schmid E 1940 Variationstatistische Untersuchungen am Gebiss pleistozaner und rezenter Leoparden und anderer Feliden Zeitschrift fur Saugetierkunde 15 1 179 Marciszak A amp Stefaniak K 2010 Two forms of cave lion Middle Pleistocene Panthera spelaea fossilis Reichenau 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss 1810 from the Bisnik Cave Poland Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 258 3 339 351 doi 10 1127 0077 7749 2010 0117 Diedrich C G 2013 Late Pleistocene leopards across Europe northernmost European German population highest elevated records in the Swiss Alps complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art Quaternary Science Reviews 76 167 193 Bibcode 2013QSRv 76 167D doi 10 1016 j quascirev 2013 05 009 Izawa M amp Nakanishi N 2015 Felidae In Ohdachi S D Ishibashi Y Iwasa M A amp Saitoh T eds The Wild Mammals of Japan Second ed Kyoto Shoukadoh Book Sellers and the Mammalogical Society of Japan pp 226 231 ISBN 978 4 87974 691 7 Chi T C Gan Y Yang T R amp Chang C H 2021 First report of leopard fossils from a limestone cave in Kenting area southern Taiwan PeerJ 9 e12020 doi 10 7717 peerj 12020 PMC 8388558 PMID 34513335 Kawata K 2001 Zoological gardens of Japan In Kisling V N ed Zoo and Aquarium History Ancient Animal Collections to Zoological Gardens Boca Raton Florida CRC Press pp 295 329 ISBN 978 0 8493 2100 9 Pirie T J Thomas R L amp Fellowes M D E 2017 Increasing game prices may alter farmers behaviours towards leopards Panthera pardus and other carnivores in South Africa PeerJ 5 e3369 doi 10 7717 peerj 3369 PMC 5452990 PMID 28584709 Spalton J A amp Al Hikmani H M 2006 The Leopard in the Arabian Peninsula Distribution and Subspecies Status PDF Cat News Special Issue 1 4 8 Archived PDF from the original on 2011 05 23 Judas J Paillat P Khoja A amp Boug A 2006 Status of the Arabian leopard in Saudi Arabia PDF Cat News Special Issue 1 11 19 Archived PDF from the original on 2015 09 19 Al Jumaily M Mallon D P Nasher A K amp Thowabeh N 2006 Status Report on Arabian Leopard in Yemen Cat News Special Issue 1 20 25 Soultan A Attum O Hamada A Hatab E B Ahmed S E Eisa A Al Sharif I Nagy A amp Shohdi W 2017 Recent observation for leopard Panthera pardus in Egypt Mammalia 81 1 115 117 doi 10 1515 mammalia 2015 0089 S2CID 90676105 Gavashelishvili A amp Lukarevskiy V 2008 Modelling the habitat requirements of leopard Panthera pardus in west and central Asia Journal of Applied Ecology 45 2 579 588 doi 10 1111 j 1365 2664 2007 01432 x Jhala Y V Qureshi Q amp Yadav S P 2020 Status of leopards in India 2018 Technical Report TR 2020 16 Report New Delhi and Dehradun National Tiger Conservation Authority Government of India and Wildlife Institute of India Arthreya V 2012 Living with Leopards Outside Protected Areas in India Conservation India Thapa K Pradhan N M B Berker J Dhakal M Bhandari A R Gurung G S Rai D P Thapa G J Shrestha S amp Singh G R 2013 High elevation record of a leopard cat in the Kangchenjunga Conservation Area Nepal Cat News 58 26 27 Kittle A M Watson A C Chanaka Kumara P H amp Nimalka Sanjeewani H K 2014 Status and distribution of the leopard in the central hills of Sri Lanka Cat News 56 28 31 Kittle A M Watson A C Kumara P H S C Sandanayake S D K C Sanjeewani H K N amp Fernando T S P 2014 Notes on the diet and habitat selection of the Sri Lankan Leopard Panthera pardus kotiya Mammalia Felidae in the central highlands of Sri Lanka Journal of Threatened Taxa 6 9 6214 6221 doi 10 11609 JoTT o3731 6214 21 Saw Sha Bwe Moo Froese G Z L amp Gray T N E 2017 First structured camera trap surveys in Karen State Myanmar reveal high diversity of globally threatened mammals Oryx 52 3 537 543 doi 10 1017 S0030605316001113 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leopard Panthera pardus delacouri in the Eastern Plains Landscape Cambodia PDF Natural History Bulletin of the Siam Society 59 39 47 Archived PDF from the original on 2016 02 22 Li S Wang D Lu Z amp Mc Shea W J 2010 Cats living with pandas The status of wild felids within giant panda range China Cat News 52 20 23 Wibisono H T Wahyudi H A Wilianto E Pinondang I M R Primajati M Liswanto D amp Linkie M 2018 Identifying priority conservation landscapes and actions for the Critically Endangered Javan leopard in Indonesia Conserving the last large carnivore in Java Island PLOS ONE 13 6 e0198369 Bibcode 2018PLoSO 1398369W doi 10 1371 journal pone 0198369 PMC 6021038 PMID 29949588 a b c d e Sunquist M E amp Sunquist F 2002 Leopard Panthera pardus Wild Cats of the World Chicago University of Chicago Press pp 318 342 ISBN 978 0 226 77999 7 Leyhausen P 1979 Cat behavior the predatory and social behavior of domestic and wild cats Berlin Garland Publishing Incorporated p 281 ISBN 9780824070175 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b c d Bailey T N 1993 The African leopard a study of the ecology and behaviour of a solitary felid New York Columbia University Press ISBN 978 1 932846 11 9 a b c Hunter L Henschel P amp Ray J C 2013 Panthera pardus Leopard In Kingdon J Happold D Butynski T Hoffmann M Happold M amp Kalina J eds Mammals of Africa London Bloomsbury Publishing pp 159 168 ISBN 978 1 4081 8996 2 Pirie T J Thomas R L Reilly B K amp Fellowes M D E 2014 Social interactions between a male leopard Panthera pardus and two generations of his offspring African Journal of Ecology 52 4 574 576 doi 10 1111 aje 12154 Jenny D 1996 Spatial organization of leopards Panthera pardus in Tai National Park Ivory Coast Is rainforest habitat a tropical haven Journal of Zoology 240 3 427 440 doi 10 1111 j 1469 7998 1996 tb05296 x Marker L L amp Dickman A J 2005 Factors affecting leopard Panthera pardus spatial ecology with particular reference to Namibian farmlands PDF South African Journal of Wildlife Research 35 2 105 115 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978 0 395 32214 7 Thinley P Rajaratnam R Lassoie J P Morreale S J Curtis P D Vernes K Leki Leki Phuntsho S Dorji T amp Dorji P 2018 The ecological benefit of tigers Panthera tigris to farmers in reducing crop and livestock losses in the eastern Himalayas Implications for conservation of large apex predators Biological Conservation 219 119 125 doi 10 1016 j biocon 2018 08 007 Karanth U K amp Sunquist M E 2000 Behavioural correlates of predation by tiger Panthera tigris leopard Panthera pardus and dhole Cuon alpinus in Nagarahole India Journal of Zoology 250 2 255 265 doi 10 1111 j 1469 7998 2000 tb01076 x Bhatnagar C Mahur M 2010 Observations on feeding behavior of a wild population of marsh crocodile in Baghdarrah Lake Udaipur Rajasthan Reptile Rap 10 16 18 Gower D Garrett K amp Stafford P 2012 Snakes Firefly Books p 60 ISBN 978 1 55407 802 8 Kiffner C Ndibalema V amp Kioko J 2012 Leopard Panthera pardus aggregation and interactions with Olive baboons Papio anubis in Serengeti National Park Tanzania African Journal of Ecology 51 1 168 171 doi 10 1111 aje 12002 a b c Nowell Kristin Jackson Peter Group IUCN SSC Cat Specialist 1996 Wild Cats Status Survey and Conservation Action Plan IUCN p 26 ISBN 978 2 8317 0045 8 Sadleir R 1966 Notes on the Reproduction of the larger Felidae International Zoo Yearbook 6 184 187 doi 10 1111 j 1748 1090 1966 tb01746 x Pacifici M Santini L Di Marco M Baisero D Francucci L Grottolo Marasini G Visconti P amp Rondinini C 2013 Generation length for mammals Nature Conservation 5 87 94 Hemmer H 1976 Gestation period and postnatal development in felids In Eaton R L ed The world s cats Vol 3 Carnivore Research Institute Univ Washington Seattle pp 143 165 Eaton R L 1977 Reproductive biology of the leopard Zoologischer Garten 47 5 329 351 a b c Kingdon Jonathan Happold David Butynski Thomas Hoffmann Michael Happold Meredith Kalina Jan 2013 05 23 Mammals of Africa A amp C Black p 166 ISBN 978 1 4081 8996 2 Leopard Panthera pardus Physical characteristics and distribution Comparative Mammalian Brain Collections Salisbury S 2014 Roxanne oldest spotted leopard in captivity dies at Acreage preserve The Palm Beach Post Archived from the original on 2014 08 11 Jacobson A P Gerngross P Lemeris J R Jr Schoonover R F Anco C Breitenmoser Wursten C Durant S M Farhadinia M S Henschel P Kamler J F Laguardia A Rostro Garcia S Stein A B amp Dollar L 2016 Leopard Panthera pardus status distribution and the research efforts across its range PeerJ 4 e1974 doi 10 7717 peerj 1974 PMC 4861552 PMID 27168983 Williams S T Williams K S Lewis B P amp Hill R A 2017 Population dynamics and threats to an apex predator outside protected areas implications for carnivore management Royal Society Open Science 4 4 161090 Bibcode 2017RSOS 461090W doi 10 1098 rsos 161090 PMC 5414262 PMID 28484625 Raza R H Chauhan D S Pasha M K S amp Sinha S 2012 Illuminating the blind spot A study on illegal trade in Leopard parts in India 2001 2010 PDF Report New Delhi TRAFFIC India WWF India Archived PDF from the original on 2020 09 24 Bergin D amp Nijman V 2014 Open Unregulated Trade in Wildlife in Morocco s Markets TRAFFIC Bulletin 26 1 65 70 Bergin D amp Nijman V 2015 Potential benefits of impending Moroccan wildlife trade laws a case study in carnivore skins Biodiversity and Conservation 25 1 199 201 doi 10 1007 s10531 015 1042 1 S2CID 34533018 Abischer T Ibrahim T Hickisch R Furrer R D Leuenberger C amp Wegmann D 2020 Apex predators decline after an influx of pastoralists in former Central African Republic hunting zones PDF Biological Conservation 241 108326 doi 10 1016 j biocon 2019 108326 S2CID 213766740 Archived PDF from the original on 2020 10 03 Gomez L amp Shepherd C R 2021 The illegal exploitation of the Javan Leopard Panthera pardus melas and Sunda Clouded Leopard Neofelis diardi in Indonesia Nature Conservation 43 43 25 39 doi 10 3897 natureconservation 43 59399 S2CID 233286106 Murphey R 1951 The Decline of North Africa Since the Roman Occupation Climatic or Human PDF Annals of the Association of American Geographers XLI 2 116 132 doi 10 1080 00045605109352048 Archived PDF from the original on 2006 09 14 a b c d e Morris D 2014 Leopard Reaktion Books pp 23 24 31 33 62 99 102 111 ISBN 9781780233185 a b Benin an African kingdom PDF London British Museum Archived PDF from the original on 2008 08 05 Retrieved 2016 03 29 Kipling R 1902 How the Leopard Got His Spots PDF Just So Stories Macmillan Haist M 1999 The Lion bloodline and kingship In Hassig D ed The Mark of the Beast The Medieval Bestiary in Art Life and Literature London Taylor amp Francis pp 3 16 ISBN 978 0 8153 2952 7 Pedersen C F 1971 The International Flag Book in Color Morrow Corbett J 1955 The Temple Tiger and More Man eaters of Kumaon Oxford Oxford University Press Anderson K 1954 The Spotted Devil of Gummalapur Nine Man Eaters and one Rogue London George Allen amp Unwin pp 36 51 Owen J 2005 Medieval Lion Skulls Reveal Secrets of Tower of London Zoo National Geographic Magazine Retrieved 2007 09 05 Further readingAllsen Thomas T 2007 Natural History and Cultural History The Circulation of Hunting Leopards in Eurasia Seventh Seventeenth Centuries In Mair Victor H ed Contact and Exchange in the Ancient World Honolulu University of Hawai i Press ISBN 978 0 8248 2884 4 DeRuiter D J Berger L R 2000 Leopards as Taphonomic Agents in dolomitic Caves Implications for bone Accumulations in the Hominid bearing Deposits of South Africa Journal of Archaeological Science 27 8 665 684 doi 10 1006 jasc 1999 0470 Schaller G B 1972 The Serengeti Lion Chicago University of Chicago Press ISBN 978 0 226 73639 6 Sanei A 2007 Analysis of leopard Panthera pardus status in Iran in Persian Tehran Sepehr Publication Center ISBN 978 964 6123 74 8 Sanei A Zakaria M Yusof E Roslan M 2011 Estimation of leopard population size in a secondary forest within Malaysia s capital agglomeration using unsupervised classification of pugmarks PDF Tropical Ecology 52 1 209 217 Archived PDF from the original on 2011 10 02 Taylor P Barrientos S Dolan C 2005 Beyond Conservation A Wildland Strategy Earthscan ISBN 978 1 84407 197 5 Zakaria M Sanei A 2011 Conservation and management prospects of the Persian and Malayan leopards Asia Life Sciences Supplement 7 1 5 External links Wikimedia Commons has media related to Panthera pardus category IUCN SSC Cat Specialist Group Panthera pardus in Africa and Panthera pardus in Asia Leopard Encyclopaedia Britannica 11th ed 1911 Retrieved from https en wikipedia org w index php title Leopard amp oldid 1154792373, wikipedia, wiki, book, books, library,

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