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The leopard (''Panthera pardus'') is one of the five extant
species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individu ...

species
in the
genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying gr ...
''
Panthera ''Panthera'' is a genus within the family (biology), family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as co ...

Panthera
'', a member of the cat
family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or social territory, typically subject to the same Politic ...
,
Felidae Felidae () is a Family (biology), family of mammals in the Order (biology), order Carnivora, colloquially referred to as cats, and constitutes a clade. A member of this family is also called a felid (). The term "cat" refers both to felids in ge ...
. It occurs in a wide range in
sub-Saharan Africa Sub-Saharan Africa (commonly called Black Africa) is, geographically, the area of the continent of Africa that lies south of the Sahara. According to the United Nations, it consists of all list of sovereign states and dependent territories i ...

sub-Saharan Africa
, in some parts of
Western Western may refer to: Places *Western, Nebraska, a village in the US *Western, New York, a town in the US *Western Creek, Tasmania, a locality in Australia *Western Junction, Tasmania, a locality in Australia *Western world, countries that ide ...

Western
and
Central Asia Central Asia is a region in Asia which stretches from the Caspian Sea in the west to China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north, including the former Soviet Union, Soviet republics of the Sov ...

Central Asia
,
Southern Russia Southern Russia or the South of Russia (russian: Юг России, ''Yug Rossii'') is a colloquial Colloquialism or colloquial language is the style (sociolinguistics), linguistic style used for casual (informal) communication. It is the most ...
, and on the Indian subcontinent to
Southeast The points of the compass are the vectors by which planet A planet is an astronomical body orbiting a star or Stellar evolution#Stellar remnants, stellar remnant that is massive enough to be Hydrostatic equilibrium, rounded by its own gravity ...

Southeast
and
East Asia East Asia is the eastern region of Asia, which is defined in both Geography, geographical and culture, ethno-cultural terms. The modern State (polity), states of East Asia include China, Japan, Mongolia, North Korea, South Korea, and Taiwan. ...

East Asia
. It is listed as
Vulnerable Vulnerable may refer to: General *Vulnerability *Vulnerability (computing) *Vulnerable adult *Vulnerable species Music Albums *Vulnerable (Marvin Gaye album), ''Vulnerable'' (Marvin Gaye album), 1997 *Vulnerable (Tricky album), ''Vulnerable'' (T ...
on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species (also known as the IUCN Red List or Red Data Book), founded in 1964, is the world's most comprehensive inventory of the global Conservation movement, conser ...
because leopard populations are threatened by
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat Ibex in an alpine habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are pr ...
and fragmentation, and are declining in large parts of the global range. The leopard is considered
locally extinct Local extinction, also known as extirpation, is the condition of a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A specie ...
in
Hong Kong Hong Kong (; , ), officially the Hong Kong Special Administrative Region of the People's Republic of China (HKSAR), is a city A city is a large human settlement.Goodall, B. (1987) ''The Penguin Dictionary of Human Geography''. London: Pe ...

Hong Kong
,
Singapore Singapore (), officially the Republic of Singapore, is a sovereign state, sovereign island city-state in maritime Southeast Asia. It lies about one degree of latitude () north of the equator, off the southern tip of the Malay Peninsula, bor ...

Singapore
,
South Korea South Korea, officially the Republic of Korea (ROK), is a country in East Asia, constituting the southern part of the Korea, Korean Peninsula and sharing a Korean Demilitarized Zone, land border with North Korea. Its western border is for ...

South Korea
,
Jordan Jordan ( ar, الأردن; tr. ' ), officially the Hashemite Kingdom of Jordan,; tr. ') is a country in Western Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion A subregion is a part of a larger region In ge ...

Jordan
,
Morocco ) , image_map = Morocco (orthographic projection, WS claimed).svg , map_caption = Location of Morocco in northwest Africa.Dark green: Undisputed territory of Morocco.Lighter green: Western Sahara, a United Nations lis ...

Morocco
,
Togo Togo (), officially the Togolese Republic (french: République togolaise), is a country in West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 17 countries of Benin ...

Togo
, the
United Arab Emirates The United Arab Emirates (UAE; ar, الإمارات العربية المتحدة ) or the Emirates ( ar, الإمارات ), is a country in Western Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion A subregio ...

United Arab Emirates
,
Uzbekistan Uzbekistan (, ; uz, Ozbekiston, ), officially the Republic of Uzbekistan ( uz, Ozbekiston Respublikasi), is a landlocked country A country is a distinct territory, territorial body or political entity. It is often referred to as the land ...

Uzbekistan
,
Lebanon Lebanon ( , ar, لُبْنَان, translit=lubnān, ), officially the Republic of Lebanon or the Lebanese Republic, is a country in Western Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion A subregion is a part ...

Lebanon
,
Mauritania Mauritania (; ar, موريتانيا, ', french: Mauritanie; Berber languages, Berber: ''Agawej'' or ''Cengit''; Pulaar language, Pulaar: ''Moritani''; Wolof language, Wolof: ''Gànnaar''; Soninke language, Soninke: ''Murutaane''), officially ...

Mauritania
,
Kuwait Kuwait (; ar, الكويت ', or ), officially the State of Kuwait ( ar, دولة الكويت '), is a country in Western Asia Western Asia, West Asia, or Southwest Asia, is the westernmost subregion A subregion is a part of a larger regi ...

Kuwait
,
Syria Syria ( ar, سُورِيَا or ar, سُورِيَة, ''Sūriyā''), officially the Syrian Arab Republic ( ar, ٱلْجُمْهُورِيَّةُ ٱلْعَرَبِيَّةُ ٱلسُّورِيَّةُ, al-Jumhūrīyah al-ʻArabīyah as-S ...

Syria
,
Libya Libya (; ar, ليبيا, Lībiyā), officially the State of Libya ( ar, دولة ليبيا, Dawlat Lībiyā), is a country in the Maghreb region in North Africa bordered by the Mediterranean Sea to the north, Egypt to Egypt–Libya border, th ...

Libya
,
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa Africa is the world's second-largest and second-most populous , after in both cases. At about 30.3 million km2 (11. ...

Tunisia
and most likely in
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia, constituting the northern part of the Korean Peninsula. It borders China and Russia to the north, at the Yalu River, Yalu (Amnok) and Tu ...

North Korea
,
Gambia The Gambia (), officially the Republic of The Gambia, is a country in West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 17 countries of Benin, Burkina Faso, Ca ...

Gambia
,
Laos , national_anthem = "Pheng Xat Lao "Pheng Sat Lāo" () is the national anthem A national anthem is a song that officially symbolizes a country A country is a distinct territory, territorial body or political entity. It is often r ...

Laos
,
Lesotho Lesotho ( , ), officially the Kingdom of Lesotho ( st, Naha ea Lesotho), is an enclaved country surrounded entirely by South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. Wi ...

Lesotho
,
Tajikistan ) , image_map = Tajikistan (orthographic projection).svg , map_caption = , capital = Dushanbe Dushanbe ( tg, Душанбе, ; ; russian: Душанбе) is the Capital city, capital and largest ...

Tajikistan
,
Vietnam Vietnam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,, group="n" is a country in Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia or SEA, is the ...

Vietnam
and
Israel Israel (; he, יִשְׂרָאֵל, translit=Yīsrāʾēl; ar, إِسْرَائِيل, translit=ʾIsrāʾīl), officially the State of Israel ( he, מְדִינַת יִשְׂרָאֵל, label=none, translit=Medīnat Yīsrāʾēl; ), is a ...

Israel
. Contemporary records suggest that the leopard occurs in only 25% of its historical global range. Compared to other wild cats, the leopard has relatively short legs and a long body with a large skull. Its fur is marked with rosettes. It is similar in appearance to the
jaguar The jaguar (''Panthera onca'') is a large cat The cat (''Felis catus'') is a domestic species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as ...

jaguar
(''Panthera onca''), but has a smaller, lighter physique, and its rosettes are generally smaller, more densely packed and without central spots. Both leopards and jaguars that are
melanistic The term melanism refers to black pigment and is derived from the gr, μελανός. Melanism is the increased development of the dark-colored pigment A pigment is a colored material that is completely or nearly insoluble in water. In contr ...
are known as
black panther A black panther is the melanistic colour variant of the leopard The leopard (''Panthera pardus'') is one of the five extant species In biology, a species is the basic unit of biological classification, classification and a taxonomic ...

black panther
s. The leopard is distinguished by its well-camouflaged fur,
opportunistic Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term c ...
hunting behaviour, broad diet, strength, and its ability to adapt to a variety of habitats ranging from
rainforest Rainforests are characterized by a closed and continuous tree canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapevine Religi ...

rainforest
to steppe, including arid and montane areas. It can run at speeds of up to . The earliest known leopard
fossil A fossil (from Classical Latin Classical Latin is the form of Latin language Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication used by humans, inc ...

fossil
s excavated in
Europe Europe is a continent A continent is any of several large landmass A landmass, or land mass, is a large region In geography Geography (from Greek: , ''geographia'', literally "earth description") is a field of scienc ...

Europe
are estimated 600,000 years old, dating to the late
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest (or lowest) division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently ...
. Leopard fossils were also found in
Japan Japan ( ja, 日本, or , and formally ) is an island country An island country or an island nation is a country A country is a distinct territory, territorial body or political entity. It is often referred to as the land of an in ...

Japan
and
Sumatra Sumatra is one of the Sunda Islands The Sunda Islands are a group of islands in the Malay Archipelago. They consist of the Greater Sunda Islands The Greater Sunda Islands are four tropical islands situated within Southeast Asia S ...

Sumatra
.


Etymology

The English name 'leopard' comes from fro, leupart or frm, liepart, that derives from la, leopardus and grc, λέοπάρδος (''leopardos''). ''Leopardos'' could be a
compound Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive structu ...
of (''leōn''), meaning lion, and (''pardos''), meaning spotted. The word ''λέοπάρδος'' originally referred to a
cheetah The cheetah (''Acinonyx jubatus'') is a large Felidae, cat native to Africa and central Iran. It is the Fastest animals, fastest land animal, estimated to be capable of running at with the fastest reliably recorded speeds being , and as suc ...

cheetah
(''Acinonyx jubatus''). 'Panther' is another common name, derived from la, panther and grc, πάνθηρ (''pánthēr''); The generic name ''Panthera'' originates in la, panthera, which refers to a hunting net for catching wild beasts that were used by the
Roman Roman or Romans most often refers to: *, the capital city of Italy *, Roman civilization from 8th century BC to 5th century AD *, the people of ancient Rome *', shortened to ''Romans'', a letter in the New Testament of the Christian Bible Roman ...
s in combats. The phonetically similar sa, पाण्डर (''pând-ara'') means pale yellow, whitish or white. The
specific nameSpecific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In Taxonomy (biology), taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), ...
la, pardus refers to a male panther. The word ''pardus'' is thought to originate from grc, παρδάλωτός (''pardalotόs'') meaning spotted like a leopard.


Characteristics

The leopard's fur is generally soft and thick, notably softer on the belly than on the back. Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in rosettes. Its belly is whitish and its ringed tail is shorter than its body. Its pupils are round. 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. 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 Rainforests are forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing and ecological func ...

rain forest
habitats. The pattern of the rosettes is unique in each individual. This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard The leopard (''Pan ...

camouflage
. Its white-tipped tail is about long, white underneath and with spots that form incomplete bands toward the tail's end. The guard hairs protecting the basal hairs are short, in face and head, and increase in length toward the flanks and the belly to about . Juveniles have woolly fur, and appear to be dark-coloured due to the densely arranged spots. Its fur tends to grow longer in colder climates. The leopard's rosettes differ from those of the
jaguar The jaguar (''Panthera onca'') is a large cat The cat (''Felis catus'') is a domestic species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as ...

jaguar
(''Panthera onca''), which are darker and with smaller spots inside. The leopard has a
diploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Sets of chromosomes refer to the number of mate ...
chromosome A chromosome is a long DNA molecule with part or all of the genome, genetic material of an organism. Most eukaryotic chromosomes include packaging proteins called histones which, aided by Chaperone (protein), chaperone proteins, bind to and ...

chromosome
number of 38. The chromosomes include four
acrocentric In this diagram of a duplicated chromosome, (2) identifies the centromere—the region that joins the two sister chromatids, or each half of the chromosome. In prophase of mitosis, specialized regions on centromeres called kinetochores attach chr ...
, five
metacentric Metacentric may refer to: * Metacentric height, the distance between the center of gravity of a ship and its metacenter * Centromere#Metacentric, Metacentric centromere, the position of a centromere on a chromatid {{disambiguation ...
, seven
submetacentric In this diagram of a duplicated chromosome, (2) identifies the centromere—the region that joins the two sister chromatids, or each half of the chromosome. In prophase of mitosis, specialized regions on centromeres called kinetochores attach chr ...
and two
telocentric In this diagram of a duplicated chromosome, (2) identifies the centromere—the region that joins the two sister chromatids, or each half of the chromosome. In prophase of mitosis, specialized regions on centromeres called kinetochores attach chr ...
pairs.


Size and weight

The leopard is
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same species exhibit different characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most animals and some plants. Differences may in ...
with males larger and heavier than females. It is muscular, with relatively short limbs and a broad head. Males stand at the shoulder, while females are tall. The head-and-body length ranges between with a long tail. Sizes vary geographically. Males weigh , and females . Some leopards in North Africa allegedly were as large as
Barbary lion The Barbary lion is an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies the Life#Bi ...

Barbary lion
s (''Panthera leo leo''). In 1913, an Algerian newspaper reported a leopard killed that allegedly measured about in total length. The maximum weight of a wild leopard in Southern Africa was about . It measured . An Indian leopard killed in
Himachal Pradesh Himachal Pradesh (; ; "Province of the Snow-laden Mountains") is a state State may refer to: Arts, entertainment, and media Literature * ''State Magazine'', a monthly magazine published by the U.S. Department of State * The State (newspaper ...

Himachal Pradesh
in 2016 measured with an estimated weight of ; it was perhaps the largest known wild leopard in India. The largest skull of a leopard was recorded in India in 1920 and measured in
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is neces ...
length, in breadth, and weighed . The skull of an African leopard measured in basal length, and in breadth, and weighed .


Variant colouration

Melanistic The term melanism refers to black pigment and is derived from the gr, μελανός. Melanism is the increased development of the dark-colored pigment A pigment is a colored material that is completely or nearly insoluble in water. In contr ...
leopards are also called
black panther A black panther is the melanistic colour variant of the leopard The leopard (''Panthera pardus'') is one of the five extant species In biology, a species is the basic unit of biological classification, classification and a taxonomic ...

black panther
s. Melanism in leopards is caused by a
recessive allele In genetics Genetics is a branch of biology concerned with the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) Though heredity had been observed for millennia, Gregor Mendel, Moravia, Moravian scientis ...
and inherited as a
recessive trait In genetics Genetics is a branch of biology concerned with the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) Though heredity had been observed for millennia, Gregor Mendel, Moravia, Moravian scientis ...
. Interbreeding in melanistic leopards produces a significantly smaller litter size than is produced by normal pairings. The black leopard is common foremost in
tropical The tropics are the region of Earth surrounding the Equator. They are delimited in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S; these latitudes correspond to ...

tropical
and subtropical moist forests like the
equatorial rainforest Equatorial generally means "of or related to an equator The Earth's equator is an imaginary planetary line that is about long in circumference. The equator divides the planet into the Northern Hemisphere and Southern Hemisphere and is loca ...
of the
Malay Peninsula The Malay Peninsula (Malay Malay may refer to: Languages * Malay language or Bahasa Melayu, a major Austronesian language spoken in Indonesia, Malaysia, Brunei and Singapore ** History of the Malay language#Old Malay, the Malay language f ...
and the
tropical rainforest Tropical rainforests are rainforest Rainforests are characterized by a closed and continuous tree canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape ...

tropical rainforest
on the slopes of some African mountains such as
Mount Kenya Mount Kenya is the highest mountain in Kenya ) , national_anthem = "Ee Mungu Nguvu Yetu "Ee Mungu Nguvu Yetu" (, ) is the national anthem of Kenya. History "Ee Mungu Nguvu Yetu"'s lyrics were originally written in Swahili la ...

Mount Kenya
. Between January 1996 and March 2009, leopards were photographed at 16 sites in the Malay Peninsula in a sampling effort of more than 1,000
camera trap A camera trap is a camera A camera is an optical Optics is the branch of physics Physics is the natural science that studies matter, its Elementary particle, fundamental constituents, its Motion (physics), motion and beh ...

camera trap
nights. Of the 445 photographs of melanistic leopards, 410 were taken in study sites south of the
Kra Isthmus The Kra Isthmus ( th, คอคอดกระ, ) in Thailand ) , royal_anthem = '' Sansoen Phra Barami''( en, "Glorify His prestige") , image_map = , map_caption = , capital = Bangkok , coordinates = , large ...
, where the non-melanistic morph was never photographed. These data indicate the near-fixation of the dark
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is one of two, or more, forms of a given gene In biology, a gene (from ''genos'' "...Wilhelm Johannsen coined the word gene to describe the Mendelian_inheritance ...
in the region. The expected time for the fixation of this recessive allele due to
genetic drift Genetic drift (allelic drift or the Sewall Wright effect) is the change in the frequency of an existing gene In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical stru ...

genetic drift
alone ranged from about 1,100 years to about 100,000 years. Pseudomelanistic leopards have also been reported. In India, nine pale and white leopards were reported between 1905 and 1967. Leopards exhibiting
erythrism Erythrism or erythrochroism refers to an unusual reddish pigmentation of an animal's hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, ap ...
were recorded between 1990 and 2015 in South Africa's Madikwe Game Reserve and in
Mpumalanga Mpumalanga () is a province A province is almost always an administrative division Administrative division, administrative unitArticle 3(1). , country subdivision, administrative region, subnational entity, first-level subdivision, ...

Mpumalanga
. The cause of this
morph Morph may refer to: Astronomy * Morphs collaboration, a collaboration that studied the evolution of spiral galaxies using the Magellan and the Hubble Space Telescope Biology * Morph (zoology), a visual or behavioral difference between organi ...
, known as a "strawberry leopard" or "pink panther", is not well understood.


Taxonomy

''Felis pardus'' was the
scientific name In taxonomy Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only ...
proposed by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement Ennoblement is the conferring of nobility—the induction of an individual into the noble social class, class. Currently only a few kingdoms still grant nob ...

Carl Linnaeus
in 1758. The generic name ''
Panthera ''Panthera'' is a genus within the family (biology), family Felidae that was named and described by Lorenz Oken in 1816 who placed all the spotted cats in this group. Reginald Innes Pocock revised the classification of this genus in 1916 as co ...

Panthera
'' was first used by
Lorenz Oken Lorenz Oken (1 August 1779 – 11 August 1851) was a German people, German natural history, naturalist, botany, botanist, biologist, and ornithology, ornithologist. Oken was born Lorenz Okenfuss (german: Okenfuß) in Bohlsbach (now part of Offenbur ...

Lorenz Oken
in 1816, who included all the known spotted cats into this group. Oken's classification was not widely accepted, and ''
Felis ''Felis'' is a genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscr ...
'' or '''' was used as the generic name until the early 20th century. The leopard was designated as the
type species In zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Convention (norm), convention in zoology that rules the formal scientific name, scientific naming of organisms treated as animals. It is al ...
of ''Panthera'' by
Joel Asaph Allen Joel Asaph Allen (July 19, 1838 – August 29, 1921) was an American zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal ...
in 1902. In 1917,
Reginald Innes Pocock Reginald Innes Pocock F.R.S. (4 March 1863 – 9 August 1947) was a British zoologist. Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edwa ...
also subordinated the
tiger The tiger (''Panthera tigris'') is the largest extant taxon, living Felidae, cat species and a member of the genus ''Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, i ...

tiger
(''P. tigris''),
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, deep-chested body, short, rounded head, round ears, and a hairy tuft at the end of its tail. It is sexually dimorphic; ...

lion
(''P. leo''), and
jaguar The jaguar (''Panthera onca'') is a large cat The cat (''Felis catus'') is a domestic species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as ...

jaguar
(''P. onca'') to ''Panthera''.


Subspecies

Following Linnaeus' first description, 27 leopard
subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interacti ...
were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered valid on the basis of
mitochondrial A mitochondrion (, plural mitochondria) is a double membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the ide ...

mitochondrial
analysis. Later analysis revealed a ninth valid subspecies, the
Arabian leopard The Arabian leopard (''Panthera pardus nimr'') is a leopard The leopard (''Panthera pardus'') is one of the five extant species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of ...
. In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa: Results of an
analysis of molecular varianceAnalysis of molecular variance (AMOVA), is a statistical model for the molecular algorithm in a single species, typically biology, biological. The name and model are inspired by ANOVA. The method was developed by Laurent Excoffier, Peter Smouse and J ...
and pairwise
fixation index The fixation index (FST) is a measure of population differentiation Human genetic variation is the genetic differences in and among populations. There may be multiple variants of any given gene in the human population (alleles), a situation calle ...
of 182 African leopard museum specimens showed that some African leopards exhibit higher genetic differences than Asian leopard subspecies.


Evolution

Results of
phylogenetic In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanism ...

phylogenetic
studies based on
nDNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA Deoxyribonucleic acid (; DNA) is a molecule File:Pentacene on Ni(111) STM.jpg, A scanning tunneling microscopy image of pentacene molecules, which consist of linear chain ...
and
mtDNA Illustration of the location of mitochondrial DNA in human cells Electron microscopy reveals mitochondrial DNA in discrete foci. Bars: 200 nm. (A) Cytoplasmic section after immunogold labelling with anti-DNA; gold particles marking mtDNA are ...

mtDNA
analysis showed that the last
common ancestor Common descent is a concept in evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biodiversity, diversity ...
of the ''Panthera'' and ''Neofelis'' genera is thought to have lived about . ''Neofelis'' diverged about from the ''Panthera'' lineage. The tiger diverged about , followed by the snow leopard about and the leopard about . The leopard is a sister taxon to a clade within ''Panthera'', consisting of the lion and the jaguar. Results of a phylogenetic analysis of chemical secretions amongst cats indicated that the leopard is closely related to the lion. 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. The leopard-lion clade diverged 3.1–1.95 million years ago. 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 Hybridization (biology), hybridized with the snow leopard at some point in their evolution. Fossils of leopard ancestors were excavated in East Africa and South Asia, dating back to the Pleistocene between 2 and . The modern leopard is suggested to have evolved in Africa about and to have Radiation (biology), radiated across Asia about 0.2 and . 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, and due to competition with the Sunda clouded leopard (''Neofelis diardi'') and the dhole (''Cuon alpinus''). 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, Puy-de-Dôme, 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 (region), Baranya in southern Hungary, Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central Poland. 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 (Baden), Mauer in Germany. Four European Pleistocene leopard subspecies were proposed. ''P. p. begoueni'' from the beginning of the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest (or lowest) division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently ...
was replaced about by ''P. p. sickenbergi'', which in turn was replaced by ''P. p. antiqua'' around 0.3 million years ago. The most recent, Panthera pardus spelaea, ''P. p. spelaea'', appeared at the beginning of the Late Pleistocene and survived until about 24,000 years ago in several parts of Europe. Leopard fossils dating to the Pleistocene were also excavated in the Japanese archipelago.


Hybrids

In 1953, a male leopard and a lioness were Crossbreeding, 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.


Distribution and habitat

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 Local extinction, extirpated in North Africa. It inhabits foremost savanna and
rainforest Rainforests are characterized by a closed and continuous tree canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapevine Religi ...

rainforest
, and areas where grasslands, woodlands, and riverine forests remain largely undisturbed. In
sub-Saharan Africa Sub-Saharan Africa (commonly called Black Africa) is, geographically, the area of the continent of Africa that lies south of the Sahara. According to the United Nations, it consists of all list of sovereign states and dependent territories i ...

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. Leopard populations on the Arabian Peninsula are small and fragmented. In southeastern Egypt, a leopard killed in 2017 was the first record in this area in 65 years. In western and central Asia, it avoids deserts, areas with long snow cover and proximity to urban centres. In the Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other ''Panthera'' species. 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 in the Shivalik Hills and Gangetic plains, Central India and Eastern Ghats, Western Ghats, the Brahmaputra River basin and hills in Northeast India. 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. In Nepal's Kanchenjunga Conservation Area, a melanistic leopard was photographed at an elevation of by a camera trap in May 2012. In Sri Lanka, leopards were recorded in Yala National Park and in unprotected forest patches, tea estates, grasslands, home gardens, Pinus, pine and eucalyptus plantations. In Myanmar, leopards were recorded for the first time by camera traps in the hill forests of Myanmar's Karen State. The Northern Tenasserim Hills, Tenasserim Forest Complex in southern Myanmar is considered a leopard stronghold. In Thailand, leopards are present in the Western Forest Complex, Kaeng Krachan National Park, Kaeng Krachan-Kui Buri National Park, Kui Buri, Khlong Saeng Wildlife Sanctuary, Khlong Saeng-Khao Sok National Park, Khao Sok protected area complexes and in Hala Bala Wildlife Sanctuary bordering Malaysia. In Peninsular Malaysia, leopards are present in Belum-Temengor, Taman Negara National Park, Taman Negara and Endau-Rompin National Parks. In Laos, leopards were recorded in Nam Et-Phou Louey National Biodiversity Conservation Area and Nam Kan National Protected Area. In Cambodia, leopards inhabit deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary and Mondulkiri Protected Forest. In southern China, leopards were recorded only in the Qinling Mountains during surveys in 11 nature reserves between 2002 and 2009. In Java, leopards inhabit dense
tropical rainforest Tropical rainforests are rainforest Rainforests are characterized by a closed and continuous tree canopy Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape ...

tropical rainforest
s and dry deciduous forests at elevations from sea level to . Outside protected areas, leopards were recorded in mixed agricultural land, secondary forest and Agroforestry, production forest between 2008 and 2014. In the Russian Far East, it inhabits temperate coniferous forests where winter temperatures reach a low of .


Behaviour and ecology

The leopard is a solitary and Territory (animal), territorial animal. 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. The roaring sequence in leopards consists mainly of grunts, also called "sawing", as it resembles the sound of sawing wood. Cubs call their mother with a ''urr-urr'' sound. The whitish spots on the back of its ears are thought to play a role in communication. 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. Leopards are active mainly from dusk till dawn and rest for most of the day and for some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking across their range at night; they may even wander up to if disturbed. In some regions, they are Nocturnality, nocturnal. In western African forests, they have been observed to be largely Diurnality, diurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons. Leopards can climb trees very skillfully, often rest on tree branches and descend from trees headfirst. They can run at over , leap over horizontally, and jump up to vertically.


Social spacing

In Kruger National Park, most leopards tend to keep apart. Males interact with their partners and cubs at times, and exceptionally this can extend beyond to two generations. Aggressive encounters are rare, typically limited to defending territories from intruders. In a South African reserve, a male was wounded in a male–male territorial battle over a carcass. 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. 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. 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. Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the Serengeti, males have home ranges of and females of ; but males in northeastern Namibia of and females of . They are even larger in arid and montane areas. In Nepal's Bardia National Park, male home ranges of and female ones of are smaller than those generally observed in Africa.


Hunting and diet

The leopard is a carnivore that prefers medium-sized prey with a body mass ranging from . 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 (animal), genet (''Genetta'' sp.) and cheetah. The largest prey killed by a leopard was reportedly a male Taurotragus, eland weighing . 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 Chinese bamboo rat, bamboo rats (''Rhizomys sinense'') and other smaller prey. The leopard depends mainly on its acute senses of hearing and vision for hunting. It primarily hunts at night in most areas. In western African forests and Tsavo National Park, they have also been observed hunting by day. They usually hunt on the ground. In the Serengeti, they have been observed to ambush prey by jumping down on it from trees. The animal stalks its prey and tries to approach as closely as possible, typically within 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 Throat clamp, by the throat and strangles them It caches kills up to apart. 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 up into a tree. It eats small prey immediately, but drags larger carcasses over several hundred metres and caches it safely in trees, bushes or even caves. 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. Average daily consumption rates of were estimated for males and of for females. 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 Schmidtia, sour grass (''Schmidtia kalahariensis'').


Enemies and competitors

In parts of its global range, the leopard is Sympatricity, sympatric with other large predators such as the
tiger The tiger (''Panthera tigris'') is the largest extant taxon, living Felidae, cat species and a member of the genus ''Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, i ...

tiger
(''Panthera tigris''),
lion The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'' native to Africa and India. It has a muscular, deep-chested body, short, rounded head, round ears, and a hairy tuft at the end of its tail. It is sexually dimorphic; ...

lion
(''P. leo''), cheetah, 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''). Leopards generally seem to avoid encounters with adult bears, but kill vulnerable bear cubs. In Sri Lanka, a few recorded vicious fights between leopards and sloth bears (''Melursus ursinus'') apparently result in both animals winding up either dead or grievously injured. While interspecies killing of full-grown leopards is generally rare, given the opportunity, both tiger and lion readily kill and consume both young and adult leopards. In the Kalahari Desert, leopards frequently lose kills to brown hyenas, if the leopard is unable to move the kill into a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills. Lions occasionally fetch leopard kills from trees. Resource partitioning occurs where leopards share their range with tigers. Leopards tend to take smaller prey, usually less than , where tigers are present. In areas where leopard and tiger are sympatric, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous. Tigers appear to inhabit the deep parts of a forest while leopards are pushed closer to the fringes. 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 prey selected, tigers and leopards seem to successfully coexist without competitive exclusion or interspecies dominance hierarchies that may be more common to the leopard's co-existence with the lion in savanna habitats. Nile crocodiles (''Crocodylus niloticus'') prey on leopards occasionally. One large adult leopard was grabbed and consumed by a large crocodile while attempting to hunt along a bank in Kruger National Park. Mugger crocodiles (''Crocodylus palustris'') reportedly killed an adult leopard in Rajasthan. An adult leopard was recovered from the stomach of a Burmese python (''Python bivittatus''). In Serengeti National Park, troops of 30–40 olive baboons (''Papio anubis'') were observed while mobbing and attacking a female leopard and her cubs.


Reproduction and life cycle

In some areas, leopards mate all year round. In Manchuria and Siberia, they mate during January and February. The female's estrous cycle lasts about 46 days, and she usually is in heat for 6–7 days. The Generation time, generation length of the leopard is 9.3 years. Gestation lasts for 90 to 105 days. Cubs are usually born in a Litter (zoology), litter of 2–4 cubs. Mortality of cubs is estimated at 41–50% during the first year. Females give birth in a cave, crevice among boulders, hollow tree or thicket. Cubs are born with closed eyes, which open four to nine days after birth. 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. 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 remain with the mother for 18–24 months. The average typical life span of a leopard is 12–17 years. The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.


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


Human interaction


Cultural significance

The leopards has 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. 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 (ruler), oba'', since the leopard was considered the king of the forest. The Ashanti people, 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. In Rudyard Kipling's ''Just So Stories'' ''How the Leopard Got His Spots'', a leopard with no spots in the Highveld, 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. 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. The leopard is a frequently used in Leopard (heraldry), heraldry, most commonly as ''Attitude_(heraldry)#Passant, passant''. The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the Lion (heraldry), heraldic lion, and the two are often used interchangeably. Naturalistic leopard-like depictions appear on the coat of arms of Benin, Coat of arms of Malawi, Malawi, Coat of arms of Somalia, Somalia, the Coat of arms of the Democratic Republic of the Congo, Democratic Republic of the Congo and Coat of arms of Gabon, Gabon, the last of which uses a black panther.


Attacks on people

The Leopard of Rudraprayag killed more than 125 people; the Panar Leopard was thought to have killed more than 400 people. Both were shot by Jim Corbett. The ''spotted devil of Gummalapur'' killed about 42 people in Karnataka, India.


In captivity

The Ancient Romans kept leopards in captivity to be slaughtered in Venatio, hunts as well as be used in Damnatio ad bestias, executions of criminals. In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities. 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 of England, Henry III by Holy Roman Emperor Frederick II, Holy Roman Emperor, Frederick II. In modern times, leopards have been Animal training, trained and Tame animal, tamed in circuses.


See also

* List of largest cats * Leopard pattern * Panther (legendary creature)


References


Further reading

* * * * * * *


External links


IUCN/SSC Cat Specialist Group: ''Panthera pardus'' in Africa
an
''Panthera pardus'' in Asia
* {{DEFAULTSORT:Leopard Panthera Felids of Asia Felids of Africa National symbols of Benin National symbols of Malawi National symbols of Somalia National symbols of the Democratic Republic of the Congo Mammals described in 1758 Articles containing video clips Big cats Taxa named by Carl Linnaeus