Leopard

• 1. Introduction

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  A leopard can drag a carcass heavier than its own body straight up into a tree, and one individual was seen hauling a young giraffe weighing nearly 125 kilograms 5.7 metres off the ground. Panthera pardus is one of five living cat species in the genus Panthera, built slender and muscular, with a pale yellowish to dark golden coat marked by dark spots grouped into rosettes. Males typically weigh 30.9 to 72 kilograms and females 20.5 to 43 kilograms, yet this mid-sized cat has the widest range of any wild cat, found across fragmented pockets of Africa and Asia. It was first described in 1758, and eight subspecies are now recognised across that range. Listed as Vulnerable on the IUCN Red List, its populations are declining in large parts of that range as habitat is lost and broken apart. How an animal this adaptable, capable of living in rainforest, steppe, desert edge and city outskirts alike, still keeps losing ground is the question running through everything that follows.

• 2. Naming And Splitting A Species

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  The English word leopard traces back through Old French leupart and Middle French liepart to the Latin leopardus and the ancient Greek leopardos. That Greek word was originally built from leon, meaning lion, and pardos, but it did not originally refer to this animal at all. Leopardos originally referred to a cheetah, Acinonyx jubatus, not to Panthera pardus. Panther, another common name for the same animal, comes from the Latin panthera and the Greek panther, and the genus name Panthera itself traces to a Latin word for a hunting net once used by the Romans to catch wild beasts for combat. Carl Linnaeus proposed the scientific name Felis pardus in 1758, and Panthera did not appear as a genus name until Lorenz Oken introduced it in 1816 to group together all of the known spotted cats. Oken's classification did not catch on right away, and Felis or Leopardus remained in use as the generic name until the early 20th century. Joel Asaph Allen designated the leopard as the type species of Panthera in 1902, and in 1917 Reginald Innes Pocock placed the tiger, lion and jaguar into the same genus alongside it. Naming the species turned out to be far simpler than counting its subspecies. Between 1794 and 1956, naturalists proposed 27 separate leopard subspecies, a number since 1996 has been cut down to just eight on the basis of mitochondrial analysis, with later work identifying a ninth valid subspecies, the Arabian leopard. In 2017, the Cat Classification Task Force confirmed eight as valid: the African leopard, the most widespread of the group though likely already gone from Mauritania, Togo, Morocco, Algeria, Tunisia and Libya; the Indian leopard, found across the Indian subcontinent, Myanmar and southern Tibet; the Javan leopard, confined to the island of Java and assessed as Endangered in 2021; the smallest subspecies, the Arabian leopard, reduced by 2023 to an estimated 100 to 120 individuals surviving in Oman and Yemen and locally extinct in Syria, Lebanon, Israel, Palestine, Jordan, Kuwait and the United Arab Emirates; a subspecies ranging from eastern Turkey and the Caucasus to the Iranian Plateau, the Hindu Kush and the western Himalayas, listed as Endangered and already gone from Uzbekistan and Tajikistan; the Amur leopard of the Russian Far East and northern China, extinct on the Korean peninsula; the Indochinese leopard, listed as Critically Endangered and already lost from Hong Kong, Singapore, Laos and Vietnam; and the Sri Lankan leopard, found only on that island and listed as Vulnerable. A separate Balochistan population, spread across southern Iran, Afghanistan and Pakistan, is cut off from populations further north by the Dasht-e Kavir and Dasht-e Lut deserts. Genetic analysis of 182 African leopard museum specimens found that some African leopard populations differ from each other more than entire Asian subspecies differ from one another.

• 3. Central Asia To Vanished Europe

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  Phylogenetic studies of nuclear and mitochondrial DNA place the geographic origin of the genus Panthera in northern Central Asia, with the leopard and lion sharing a more recent common ancestor than either shares with the tiger or snow leopard. The leopard-lion clade spread across the Asian and African Palearctic from at least the early Pliocene onward, before the two split apart between 3.1 and 1.95 million years ago. A 2016 study found that the mitochondrial genomes of leopards, lions and snow leopards resemble each other more closely than their nuclear genomes do, suggesting their ancestors interbred with the snow leopard lineage at some point. The oldest unambiguous leopard fossils come from Eastern Africa, dated to around 2 million years ago, and leopards reached Eurasia during the late Early to Middle Pleistocene, roughly 1.2 to 0.6 million years ago. Fossil bones and teeth possibly from the Pliocene turned up at Perrier in France and at Valdarno in Italy, and until 1940 similar Pleistocene-era remains were excavated at some 40 European sites, from Furninha Cave near Lisbon and the Genista Caves in Gibraltar to Santander Province in Spain, and onward to Derby in England, Přerov in the Czech Republic and the Baranya region of southern Hungary. Four European Pleistocene subspecies have been proposed in sequence, ending with P. p. spelaea, which survived until about 11,000 years ago and possibly lingered into the early Holocene on the Iberian Peninsula. Cave paintings at Chauvet Cave offer indirect evidence that leopards once lived in Europe, and further fossil remains have turned up in Biśnik Cave in Poland, the Iberian Mountains, the Italian Peninsula, the Balkans and even southern Taiwan.

• 4. Rosettes, Melanism And Size

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  A leopard's fur runs paler and softer on the belly than on the back, coloured anywhere from pale cream in arid regions to deep golden in forest and mountain habitat, always marked with dark rosettes that fade toward the white underbelly. Those rosettes differ by region and from those of the jaguar, running circular in East African populations, squarish in Southern African ones, and larger across Asian populations, while jaguar rosettes are darker with smaller spots inside them. The leopard carries a diploid chromosome number of 38. Some individuals turn out fully black, known as melanistic leopards or black panthers, a colouring caused by a recessive allele. Rarer still is erythrism, a reddish morph nicknamed the strawberry leopard or pink panther, recorded in South Africa's Madikwe Game Reserve and in Mpumalanga between 1990 and 2015, its cause still not well understood; in India, nine pale or white leopards were reported between 1905 and 1967. Males stand 60 to 70 centimetres at the shoulder and females 57 to 64 centimetres, with a head-and-body length of 92 to 183 centimetres and a tail running 66 to 102 centimetres. Males typically weigh 30.9 to 72 kilograms, occasionally reaching 91 kilograms, while females weigh 20.5 to 43 kilograms; leopards from South Africa's Cape Province run smaller still, with males reaching only 20 to 45 kilograms. The heaviest wild leopard recorded in Southern Africa weighed around 96 kilograms and measured 262 centimetres, while an Indian leopard killed in Himachal Pradesh in 2016 measured 261 centimetres and was estimated at 78.5 kilograms, possibly the largest wild leopard ever recorded in India. The largest known leopard skull, found in India in 1920, measured 28 centimetres in basal length and weighed a full kilogram.

• 5. A Range Under Pressure

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  No wild cat covers more ground than the leopard, spanning savanna, rainforest, arid country and mountain habitat across Africa and Asia, though its populations are now fragmented and shrinking. Its range in West Africa has fallen by an estimated 95 percent, and by 97 percent across the Sahara, even as it remains numerous in parts of sub-Saharan Africa where other large cats have already vanished. In southeastern Egypt, a leopard found killed in 2017 marked the first confirmed sighting there in 65 years. In India's tiger range landscapes, the forest-dwelling leopard population was estimated in 2020 at between 12,172 and 13,535 individuals, surveyed from the Shivalik Hills and Gangetic plains to Central India, the Eastern and Western Ghats, and the hills of Northeast India; a melanistic leopard was even photographed at 4300 metres elevation in Nepal's Kanchenjunga Conservation Area in 2012. Pakistan's Galyat region held an estimated 18 to 24 individuals in 2017, within a broader northern tract of Pakistan measuring 4543 square kilometres, which holds a notably high population density. Camera traps recorded the leopard for the first time in Myanmar's Karen State hill forests, and it persists in Thailand's Western Forest Complex, in Laos, in Cambodia's Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest, and across nature reserves in China's Qinling Mountains surveyed between 2002 and 2009. In Java, it ranges from sea level up to 2540 metres, and in the Russian Far East it endures temperate coniferous forest winters. Trophy hunting and poaching compound the loss: between 2002 and 2012, at least four leopards a week were estimated poached in India for the illegal skin and bone trade, and a 2013 survey of Moroccan cities turned up 37 leopard skins in seven weeks, rising to 43 in a follow-up survey the next year, with vendors admitting the skins had been imported from sub-Saharan Africa. In the Central African Republic's Chinko area, the population fell from 97 individuals in 2012 to 50 by 2017 as armed livestock herders moved through with poison and snares. In Java, seizures between 2011 and 2019 turned up body parts from 51 leopards, including six live animals, 12 skins, 13 skulls, 20 canines and 22 claws. Today the species occupies only about 25 percent of its historical range.

• 6. Ambush, Cache And Compete

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  A leopard stalks to within about 5 metres of its prey before pouncing, killing small animals with a bite to the back of the neck and strangling larger ones by the throat. It favours medium-sized prey weighing 10 to 40 kilograms, ungulates like impala, bushbuck, common duiker and chital above all, though more than 100 prey species have been recorded, from primates such as guenons and gray langurs to smaller carnivores like black-backed jackal and even cheetah. The largest kill ever reported was a male eland weighing 900 kilograms. Powerful jaw muscles let it drag a carcass heavier than itself into a tree, caching kills up to 2 kilometres apart to keep them from rivals; a leopard falling onto prey from a height of 2.69 metres reaches 25 kilometres per hour in 0.7 seconds, timed to avoid the horns of a struggling male. Average daily food intake runs to 3.5 kilograms for males and 2.8 for females, and in the southern Kalahari Desert leopards go without drinking for two to three days at a stretch, drawing moisture instead from prey and from plants like gemsbok cucumber and watermelon. It shares its range with formidable rivals. In Africa's predator guild alongside lions, cheetahs, hyenas and African wild dogs, the leopard outranks only the cheetah; lions cause up to 22 percent of leopard deaths in South Africa's Sabi Sand Game Reserve, while spotted hyenas are responsible for as much as 50 percent of stolen leopard kills there, and lions are skilled enough to fetch a leopard's kill straight out of a tree. In Asia, both tigers and dholes dominate leopards during direct encounters; in Nagarhole National Park, the average leopard kill weighed 37.6 kilograms against 91.5 kilograms for tigers and 43.4 for dholes, and when tiger numbers recovered in Rajaji National Park during the 2000s, leopard densities fell in response.

• 7. A Solitary Life With Cubs

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  Outside the mating season, a leopard lives and hunts alone, walking as far as 25 kilometres across its territory in a single night and up to 75 kilometres if disturbed. In Kruger National Park, most individuals keep roughly 1 kilometre of spacing between each other, and aggressive encounters stay rare, mostly limited to territorial disputes over a carcass. Home ranges vary enormously by region: males in the Serengeti hold 33 to 38 square kilometres against 14 to 16 for females, while in northeastern Namibia male ranges stretch to 451 square kilometres and female ranges to 188, shrinking again in Nepal's Bardia National Park to 48 square kilometres for males and just 5 to 7 for females. Females give birth in a cave, a crevice among boulders, a hollow tree or a thicket, to a litter of two to four cubs after a gestation of 90 to 105 days, with cub mortality estimated at 41 to 50 percent in the first year, predation being the leading cause. Newborn cubs are born with closed eyes that open four to nine days later, and males are known to kill cubs sired by rivals to bring a female back into heat. Cubs begin eating meat around nine weeks old, start following their mother on hunts near three months, and can likely fend for themselves by one year, though they typically stay with her for 18 to 24 months before separating, sometimes travelling on afterward with their littermates for months. Both sexes reach sexual maturity around two to two and a half years old, and the species has a generation length of 9.3 years and a typical lifespan of 12 to 17 years; the oldest known leopard, a captive female, lived to 24 years, 2 months and 13 days.

• 8. Kings, Spots And Spotless Lions

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  In Greek mythology, the leopard was the animal of Dionysus, who was depicted wearing its skin and being carried by leopards, and who was said in one myth to have been rescued from pirates by two of them. Roman mosaics from North African sites still show fauna that survives today only in tropical Africa. During the Benin Empire, the leopard was carved into engravings and sculptures to symbolise the power of the king, or oba, since it was considered the ruler of the forest, and the Ashanti people likewise reserved a ceremonial leopard stool for their king alone. In Rudyard Kipling's story How the Leopard Got His Spots, a spotless leopard living on the Highveld travels to the forest with his hunting partner, the Ethiopian, and the pair change their colouring together. Heraldry borrowed the leopard too, most often shown passant and, oddly, stripped of its spots and given a mane, making it nearly indistinguishable from a heraldic lion; naturalistic leopards appear on the coats of arms of Benin, Malawi, Somalia, the Democratic Republic of the Congo and Gabon, where the design uses a black panther. Two leopards became infamous man-eaters in India, both shot by the British hunter Jim Corbett: the Leopard of Rudraprayag, blamed for more than 125 deaths, and the Panar Leopard, thought responsible for over 400. In captivity, ancient Romans kept leopards to be killed in staged hunts and to execute condemned criminals, while in Benin they were paraded as mascots and offered as sacrifices to deities. A menagerie founded by King John of England once held leopards at the Tower of London, and around 1235 Holy Roman Emperor Frederick II gave three of the animals to Henry III, a gift that outlasted its own century and still marks one of the earliest recorded appearances of the species in England.