The tiger (Panthera tigris) is a large cat and a member of the genus Panthera native to Asia. It has a powerful, muscular body with a large head and paws, a long tail and orange fur with black, mostly vertical stripes. It is traditionally classified into nine recent subspecies, though some recognise only two subspecies, mainland Asian tigers and the island tigers of the Sunda Islands.
Throughout the tiger's range, it inhabits mainly forests, from coniferous and temperate broadleaf and mixed forests in the Russian Far East and Northeast China to tropical and subtropical moist broadleaf forests on the Indian subcontinent and Southeast Asia. The tiger is an apex predator and preys mainly on ungulates, which it takes by ambush. It lives a mostly solitary life and occupies home ranges, defending these from individuals of the same sex. The range of a male tiger overlaps with that of multiple females with whom he mates. Females give birth to usually two or three cubs that stay with their mother for about two years. When becoming independent, they leave their mother's home range and establish their own.
Since the early 20th century, tiger populations have lost at least 93% of their historic range and are locally extinct in West and Central Asia, in large areas of China and on the islands of Java and Bali. Today, the tiger's range is severely fragmented. It is listed as Endangered on the IUCN Red List of Threatened Species, as its range is thought to have declined by 53% to 68% since the late 1990s. Major threats to tigers are habitat destruction and fragmentation due to deforestation, poaching for fur and the illegal trade of body parts for medicinal purposes. Tigers are also victims of human–wildlife conflict as they attack and prey on livestock in areas where natural prey is scarce. The tiger is legally protected in all range countries. National conservation measures consist of action plans, anti-poaching patrols and schemes for monitoring tiger populations. In several range countries, wildlife corridors have been established and tiger reintroduction is planned.
The tiger is among the most popular of the world's charismatic megafauna. It has been kept in captivity since ancient times and has been trained to perform in circuses and other entertainment shows. The tiger featured prominently in the ancient mythology and folklore of cultures throughout its historic range and has continued to appear in culture worldwide.
Etymology
The Old English tigras derives from Old French tigre, from Latin tigris, which was a borrowing from tigris (Ancient Greek: τίγρις).[4] Since ancient times, the word tigris has been suggested to originate from the Armenian or Persian word for 'arrow', which may also be the origin of the name for the river Tigris.[5][6] However, today, the names are thought to be homonyms, and the connection between the tiger and the river is doubted.[6]
Taxonomy
In 1758, Carl Linnaeus described the tiger in his work Systema Naturae and gave it the scientific name Felis tigris, as the genus Felis was being used for all cats at the time. His scientific description was based on descriptions by earlier naturalists such as Conrad Gessner and Ulisse Aldrovandi.[2] In 1929, Reginald Innes Pocock placed the species in the genus Panthera using the scientific name Panthera tigris.[7][8]
Subspecies
Nine recent tiger subspecies have been proposed between the early 19th and early 21st centuries, namely the Bengal, Malayan, Indochinese, South China, Siberian, Caspian, Javan, Bali and Sumatran tigers.[9][10] The validity of several tiger subspecies was questioned in 1999 as most putative subspecies were distinguished on the basis of fur length and colouration, striping patterns and body size of specimens in natural history museum collections that are not necessarily representative for the entire population. It was proposed to recognise only two tiger subspecies as valid, namely P. t. tigris in mainland Asia and the smaller P. t. sondaica in the Greater Sunda Islands.[11]
This two-subspecies proposal was reaffirmed in 2015 through a comprehensive analysis of morphological, ecological and mitochondrial DNA (mtDNA) traits of all putative tiger subspecies.[10] In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group revised felid taxonomy
The cat (Felis catus), also referred to as the domestic cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the domestication of the cat occurred in the Near East around 7500 BC. It is commonly kept as a pet and farm cat, but also ranges freely as a feral cat avoiding human contact. It is valued by humans for companionship and its ability to kill vermin. Its retractable claws are adapted to killing small prey species such as mice and rats. It has a strong, flexible body, quick reflexes, and sharp teeth, and its night vision and sense of smell are well developed. It is a social species, but a solitary hunter and a crepuscular predator. Cat communication includes vocalizations—including meowing, purring, trilling, hissing, growling, and grunting—as well as body language. It can hear sounds too faint or too high in frequency for human ears, such as those made by small mammals. It secretes and perceives pheromones.
Female domestic cats can have kittens from spring to late autumn in temperate zones and throughout the year in equatorial regions, with litter sizes often ranging from two to five kittens. Domestic cats are bred and shown at events as registered pedigreed cats, a hobby known as cat fancy. Animal population control of cats may be achieved by spaying and neutering, but their proliferation and the abandonment of pets has resulted in large numbers of feral cats worldwide, contributing to the extinction of bird, mammal, and reptile species.
As of 2017, the domestic cat was the second most popular pet in the United States, with 95.6 million cats owned and around 42 million households owning at least one cat. In the United Kingdom, 26% of adults have a cat, with an estimated population of 10.9 million pet cats as of 2020. As of 2021, there were an estimated 220 million owned and 480 million stray cats in the world.
Etymology and naming
The origin of the English word cat, Old English catt, is thought to be the Late Latin word cattus, which was first used at the beginning of the 6th century.[4] The Late Latin word may be derived from an unidentified African language.[5] The Nubian word kaddîska 'wildcat' and Nobiin kadīs are possible sources or cognates.[6]
The forms might also have derived from an ancient Germanic word that was absorbed into Latin and then into Greek, Syriac, and Arabic.[7] The word may be derived from Germanic and Northern European languages, and ultimately be borrowed from Uralic, cf. Northern Sámi gáđfi, 'female stoat', and Hungarian hölgy, 'lady, female stoat'; from Proto-Uralic *käďwä, 'female (of a furred animal)'.[8]
The English puss, extended as pussy and pussycat, is attested from the 16th century and may have been introduced from Dutch poes or from Low German puuskatte, related to Swedish kattepus, or Norwegian pus, pusekatt. Similar forms exist in Lithuanian puižė and Irish puisín or puiscín. The etymology of this word is unknown, but it may have arisen from a sound used to attract a cat.[9][10]
A male cat is called a tom or tomcat[11] (or a gib,[12] if neutered). A female is called a queen[13][14] (or sometimes a molly,[15] if spayed). A juvenile cat is referred to as a kitten. In Early Modern English, the word kitten was interchangeable with the now-obsolete word catling.[16] A group of cats can be referred to as a clowder, a glaring,[17] or a colony.[18]
Taxonomy
The scientific name Felis catus was prop
It was long thought that the domestication of the cat began in ancient Egypt, where cats were venerated from around 3100 BC.[33][34] However, the earliest known indication for the taming of an African wildcat was excavated close by a human Neolithic grave in Shillourokambos, southern Cyprus, dating to about 7500–7200 BC. Since there is no evidence of native mammalian fauna on Cyprus, the inhabitants of this Neolithic village most likely brought the cat and other wild mammals to the island from the Middle Eastern mainland.[35] Scientists therefore assume that African wildcats were attracted to early human settlements in the Fertile Crescent by rodents, in particular the house mouse (Mus musculus), and were tamed by Neolithic farmers. This mutual relationship between early farmers and tamed cats lasted thousands of years. As agricultural practices spread, so did tame and domesticated cats.[32][36] Wildcats of Egypt contributed to the maternal gene pool of the domestic cat at a later time.[37]
The earliest known evidence for the occurrence of the domestic cat in Greece dates to around 1200 BC. Greek, Phoenician, Carthaginian and Etruscan traders introduced domestic cats to southern Europe.[38] By the 5th century BC, they were familiar animals around settlements in Magna Graecia and Etruria.[39] During the Roman Empire, they were introduced to Corsica and Sardinia before the beginning of the 1st century AD.[40] By the end of the Western Roman Empire in the 5th century, the Egyptian domestic cat lineage had arrived in a Baltic Sea port in northern Germany.[37]
The leopard cat (Prionailurus bengalensis) was tamed independently in China around 5500 BC. This line of partially domesticated cats leaves no trace in the domestic cat populations of today.[41]
During domestication, cats have undergone only minor changes in anatomy and behavior, and they are still capable of surviving in the wild. Several natural behaviors and characteristics of wildcats may have pre-adapted them for domestication as pets. These traits include their small size, social nature, obvious body language, love of play, and high intelligence. Since they practice rigorous grooming habits and have an instinctual drive to bury and hide their urine and feces, they are generally much less messy than other domesticated animals. Captive Leopardus cats may also display affectionate behavior toward humans but were not domesticated.[42] House cats often mate with feral cats.[43] Hybridization between domestic and other Felinae species is also possible, producing hybrids such as the Kellas cat in Scotland.[44][45]
Development of cat breeds started in the mid 19th century.[46] An analysis of the domestic cat genome revealed that the ancestral wildcat genome was significantly altered in the process of domestication, as specific mutations were selected to develop cat breeds.[47] Most breeds are founded on random-bred domestic cats. Genetic diversity of these breeds varies between regions, and is lowest in purebred populations, which show more than 20 deleterious genetic disorders.[48]
CharacteristicsMain article: Cat anatomySizeDiagram of the general anatomy of a male domestic cat
The domestic cat has a smaller skull and shorter bones than the European wildcat.[49] It averages about 46 cm (18 in) in head-to-body length and 23–25 cm (9.1–9.8 in) in height, with about 30 cm (12 in) long tails. Males are larger than females.[50] Adult domestic cats typically weigh 4–5 kg (8.8–11.0 lb).[29]
Skeleton
Cats have seven cervical vertebrae (as do most mammals); 13 thoracic vertebrae (humans have 12); seven lumbar vertebrae (humans have five); three sacral vertebrae (as do most mammals, but humans have five); and a variable number of caudal vertebrae in the tail (humans have only three to five vestigial caudal vertebrae, fused into an internal coccyx).[51]: 11 The extra lumbar and thoracic vertebrae account for the cat's spinal mobility and flexibility. Attached to the spine are 13 ribs, the shoulder, and the pelvis.[51]: 16 Unlike human arms, cat forelimbs are attached to the shoulder by free-floating clavicle bones which allow them to pass their body through any space into which they can fit their head.[52]
SkullCat skullA cat with exposed teeth and claws
Most cat
The cat is digitigrade. It walks on the toes, with the bones of the feet making up the lower part of the visible leg.[61] Unlike most mammals, it uses a "pacing" gait and moves both legs on one side of the body before the legs on the other side. It registers directly by placing each hind paw close to the track of the corresponding fore paw, minimizing noise and visible tracks. This also provides sure footing for hind paws when navigating rough terrain. As it speeds up from walking to trotting, its gait changes to a "diagonal" gait: The diagonally opposite hind and fore legs move simultaneously.[62]
osed by Carl Linnaeus in 1758 for a domestic cat.[1][2] Felis catus domesticus was proposed by Johann Christian Polycarp Erxleben in 1777.[3] Felis daemon proposed by Konstantin Satunin in 1904 was a black cat from the Transcaucasus, later identified as a domestic cat.[19][20]
In 2003, the International Commission on Zoological Nomenclature ruled that the domestic cat is a distinct species, namely Felis catus.[21][22] In 2007, the modern domesticated subspecies F. silvestris catus sampled worldwide was considered to have probably descended from the African wildcat (F. lybica), following results of phylogenetic research.[23][24][a] In 2017, the IUCN Cat Classification Taskforce followed the recommendation of the ICZN in regarding the domestic cat as a distinct species, Felis catus.[25]
in accordance with the 2015 two-subspecies proposal and recognised only P. t. tigris and P. t. sondaica.[12] Results of a 2018 whole-genome sequencing study of 32 samples from the six living putative subspecies—the Bengal, Malayan, Indochinese, South China, Siberian and Sumatran tiger—found them to be distinct and separate clades.[13] These results were corroborated in 2021 and 2023.[14][15] The Cat Specialist Group states that "Given the varied interpretations of data, the [subspecific] taxonomy of this species is currently under review by the IUCN SSC Cat Specialist Group."[16]
The following tables are based on the classification of the tiger as of 2005,[9] and also reflect the classification recognised by the Cat Classification Task Force in 2017.[12]
Panthera tigris tigris (Linnaeus, 1758)[2]PopulationDescriptionImageBengal tiger formerly P. t. tigris (Linnaeus, 1758)[2]This population inhabits the Indian subcontinent.[17] The Bengal tiger has shorter fur than tigers further north,[8] with a light tawny to orange-red colouration,[8][18] and relatively long and narrow nostrils.[19]† Caspian tiger formerly P. t. virgata (Illiger, 1815)[20]This population occurred from Turkey to around the Caspian Sea.[17] It had bright rusty-red fur with thin and closely spaced brownish stripes,[21] and a broad occipital bone.[11] Genetic analysis revealed that it was closely related to the Siberian tiger.[22] It has been extinct since the 1970s.[23]Siberian tiger formerly P. t. altaica (Temminck, 1844)[24]This population lives in the Russian Far East, Northeast China and possibly North Korea.[17] The Siberian tiger has long hair and dense fur.[24] Its ground colour varies widely from ochre-yellow in winter to more reddish and vibrant after moulting.[25] The skull is shorter and broader than the skulls of tigers further south.[19]South China tiger formerly P. t. amoyensis (Hilzheimer, 1905)[26]This tiger historically lived in south-central China.[17] The skulls of the five type specimens had shorter carnassials and molars than tigers from India, a smaller cranium, orbits set closer together and larger postorbital processes; skins were yellowish with rhombus-like stripes.[26] It has a unique mtDNA haplotype due to interbreeding with ancient tiger lineages.[12][27][28] It is extinct in the wild as there has not been a confirmed sighting since the 1970s,[1] and survives only in captivity.[15]Indochinese tiger formerly P. t. corbetti (Mazák, 1968)[29]This tiger population occurs on the Indochinese Peninsula.[17] Indochinese tiger specimens have smaller craniums than Bengal tigers and appear to have darker fur with somewhat thin stripes.[29][30]Malayan tiger formerly P. t. jacksoni (Luo et al., 2004)[31]The Malayan tiger was proposed as a distinct subspecies on the basis of mtDNA and micro-satellite sequences that differ from the Indochinese tiger.[31] It does not differ significantly in fur colour or skull size from Indochinese tigers.[30] There is no clear geographical barrier between tiger populations in northern Malaysia and southern Thailand.[1]Panthera tigris sondaica (Temminck, 1844)[12]PopulationDescriptionImage†Javan tiger formerly P. t. sondaica (Temminck, 1944)[24]This tiger was described based on an unspecified number of skins with short and smooth hair.[24] Tigers from Java were small compared to tigers of the Asian mainland, had relatively elongated skulls compared to the Sumatran tiger and longer, thinner and more numerous stripes.[30] The Javan tiger is thought to have gone extinct by the 1980s.[23]†Bali tiger formerly P. t. balica (Schwarz, 1912)[32]This tiger occurred on Bali and had brighter fur and a smaller skull than the Javan tiger.[32][33] A typical feature of Bali tiger skulls is the narrow occipital bone, which is similar to the Javan tiger's skull.[34] The tiger went extinct in the 1940s.[23]Sumatran tiger formerly P. t. sumatrae (Pocock, 1929)[35]The type specimen from Sumatra had dark fur.[35] The Sumatran tiger has particularly long hair around the face,[17] thick body stripes and a broader and smaller nasal bone than other island tigers.[30]EvolutionPanthera
Snow leopard
Tiger
Jaguar
Leopard
Lion
Phylogeny of the genus Panthera based on a 2016 nuclear DNA study[36]
The tiger shares the genus Panthera with the lion, leopard, jaguar and snow leopard. Results of genetic analyses indicate that the tiger and snow leopard are sister species whose lineages split from each other between 2.70 and 3.70 million years ago.[37] The tiger's whole genome sequencing shows repeated sequences that parallel those in other cat genomes.[38]
The fossil species Panthera palaeosinensis of early Pleistocene northern China was described as a possible tiger ancestor when it was discovered in 1924, but modern cladistics places it as basal to modern Panthera.[39][40] Panthera zdanskyi lived around the same time and place, and was suggested to be a sister species of the modern tiger when it was examined in 2014.[39] However, as of 2023, at least two subsequent studies considered P. zdanskyi likely to be a synonym of P. palaeosinensis, noting that its proposed differences from that species fell within the range of individual variation.[41][42] The earliest appearance of the modern tiger species in the fossil record are jaw fragments from Lantion in China that are dated to the early Pleistocene.[39]
Middle- to late-Pleistocene tiger fossils have been found throughout China, Sumatra and Java. Prehistoric subspecies include Panthera tigris trinilensis and P. t. soloensis of Java and Sumatra and P. t. acutidens of China; late Pleistocene and early Holocene fossils of tigers have also been found in Borneo and Palawan, Philippines.[43] Fossil specimens of tigers have also been reported from the Middle-Late Pleistocene of Japan.[44] Results of a phylogeographic study indicate that all living tigers have a common ancestor that lived between 108,000 and 72,000 years ago.[31] Genetic studies suggest that the tiger population contracted around 115,000 years ago due to glaciation. Modern tiger populations originated from a refugium in Indochina and spread across Asia after the Last Glacial Maximum. As they colonised northeastern China, the ancestors of the South China tiger intermixed with a relict tiger population.[28][27]
HybridsFurther information: Felid hybrids and Panthera hybrid
Tigers can interbreed with other Panthera cats and have done so in captivity. The liger is the offspring of a female tiger and a male lion and the tigon the offspring of a male tiger and a female lion.[45] The lion sire passes on a growth-promoting gene, but the corresponding growth-inhibiting gene from the female tiger is absent, so that ligers grow far larger than either parent species. By contrast, the male tiger does not pass on a growth-promoting gene while the lioness passes on a growth inhibiting gene; hence, tigons are around the same size as their parents.[46] Since they often develop life-threatening birth defects and can easily become obese, breeding these hybrids is regarded as unethical.[45]
CharacteristicsTiger skeleton from Royal Natural History Volume 1 (1839)
The tiger has a typical felid morphology, with a muscular body, shortened legs, strong forelimbs with wide front paws, a large head and a tail that is about half the length of the rest of its body.[47][48] It has five digits, including a dewclaw, on the front feet and four on the back, all of which have retractile claws that are compact and curved, and can reach 10 cm (3.9 in) long.[47][49] The ears are rounded and the eyes have a round pupil.[47] The snout ends in a triangular, pink tip with small black dots, the number of which increase with age.[50] The tiger's skull is robust, with a constricted front region, proportionally small, elliptical orbits, long nasal bones and a lengthened cranium with a large sagittal crest.[51][47] It resembles a lion's skull, but differs from it in the concave or flattened underside of the lower jaw and in its longer nasals.[51][43] The tiger has 30 fairly robust teeth and its somewhat curved canines are the longest in the cat family at 6.4–7.6 cm (2.5–3.0 in).[47][52]
The tiger has a head-body length of 1.4–2.8 m (4 ft 7 in – 9 ft 2 in) with a 0.6–1.1 m (2 ft 0 in – 3 ft 7 in) tail and stands 0.8–1.1 m (2 ft 7 in – 3 ft 7 in) at the shoulder.[53] The Siberian and Bengal tigers are the largest.[47] Male Bengal tigers weigh 200–260 kg (440–570 lb), and females weigh 100–160 kg (220–350 lb); island tigers are the smallest, likely due to insular dwarfism.[11] Male Sumatran tigers weigh 100–140 kg (220–310 lb), and females weigh 75–110 kg (165–243 lb).[54] The tiger is popularly thought to be the largest living felid species; but since tigers of the different subspecies and populations vary greatly in size and weight, the tiger's average size may be less than the lion's, while the largest tigers are bigger than their lion counterparts.[43]
CoatSiberian tiger coat on flank (side)
The tiger's coat usually has short hairs, reaching up to 35 mm (1.4 in), though the hairs of the northern-living Siberian tiger can reach 105 mm (4.1 in). Belly hairs tend to be longer than back hairs. The density of their fur is usually thin, though the Siberian tiger develops a particularly thick winter coat. The tiger has lines of fur around the face and long whiskers, especially in males.[47] It has an orange colouration that varies from yellowish to reddish.[55] White fur covers the underside, from head to tail, along with the inner surface of the legs and parts of the face.[47][56] On the back of the ears, it has a prominent white spot, which is surrounded by black.[47] The tiger is marked with distinctive black or dark brown stripes, which are uniquely patterned in each individual.[47][57] The stripes are mostly vertical, but those on the limbs and forehead are horizontal. They are more concentrated towards the backside and those on the trunk may reach under the belly. The tips of stripes are generally sharp and some may split up or split and fuse again. Tail stripes are thick bands and a black tip marks the end.[56]
The tiger is one of only a few striped cat species.[58] Stripes are advantageous for camouflage in vegetation with vertical patterns of light and shade, such as trees, reeds and tall grass.[57][59] This is supported by a Fourier analysis study showing that the striping patterns line up with their environment.[60] The orange colour may also aid in concealment, as the tiger's prey is colour blind and possibly perceives the tiger as green and blended in with the vegetation.[61]
Colour variationsPseudo-melanistic white tiger
The three colour variants of Bengal tigers – nearly stripeless snow-white, white and golden – are now virtually non-existent in the wild due to the reduction of wild tiger populations but continue in captive populations. The white tiger has a white background colour with sepia-brown stripes. The golden tiger is pale golden with reddish-brown stripes. The snow-white tiger is a morph with extremely faint stripes and a pale sepia-brown ringed tail. White and golden morphs are the result of an autosomal recessive trait with a white locus and a wideband locus, respectively. The snow-white variation is caused by polygenes with both white and wideband loci.[62] The breeding of white tigers is controversial, as they have no use for conservation. Only 0.001% of wild tigers have the genes for this colour morph and the overrepresentation of white tigers in captivity is the result of inbreeding. Hence, their continued breeding will risk both inbreeding depression and loss of genetic variability in captive tigers.[63]
Pseudo-melanistic tigers with thick, merged stripes have been recorded in Simlipal National Park and three Indian zoos; a population genetic analysis of Indian tiger samples revealed that this phenotype is caused by a mutation of a transmembrane aminopeptidase gene. Around 37% of the Simlipal tiger population has this feature, which has been linked to genetic isolation.[64]
Distribution and habitatCamera trap of a Siberian tiger in Russia
The tiger historically ranged from eastern Turkey, northern Iran and Afghanistan to Central Asia and from northern Pakistan through the Indian subcontinent and Indochina to southeastern Siberia, Sumatra, Java and Bali.[47] As of 2022, it inhabits less than 7% of its historical distribution and has a scattered range in the Indian subcontinent, the Indochinese Peninsula, Sumatra, northeastern China and the Russian Far East.[1] As of 2020, India had the largest extent of global tiger habitat with 300,508 km2 (116,027 sq mi), followed by Russia with 195,819 km2 (75,606 sq mi).[65]
The tiger mainly lives in forest habitats and is highly adaptable.[54] Records in Central Asia indicate that it primarily inhabited Tugay riverine forests and hilly and lowland forests in the Caucasus.[66] In the Amur-Ussuri region of Russia and China, it inhabits Korean pine and temperate broadleaf and mixed forests; riparian forests serve as dispersal corridors, providing food and water for both tigers and ungulates.[67] On the Indian subcontinent, it inhabits mainly tropical and subtropical moist broadleaf forests, temperate broadleaf and mixed forests, tropical moist evergreen forests, tropical dry forests, alluvial plains and the mangrove forests of the Sundarbans.[68] In the Eastern Himalayas, it was documented in temperate forest up to an elevation of 4,200 m (13,800 ft) in Bhutan, of 3,630 m (11,910 ft) in the Mishmi Hills and of 3,139 m (10,299 ft) in Mêdog County, southeastern Tibet.[69][70][71] In Thailand, it lives in deciduous and evergreen forests.[72] In Sumatra, it inhabits lowland peat swamp forests and rugged montane forests.[73]
Population density
Camera trapping during 2010–2015 in the deciduous and subtropical pine forest of Jim Corbett National Park, northern India revealed a stable tiger population density of 12–17 individuals per 100 km2 (39 sq mi) in an area of 521 km2 (201 sq mi).[74] In northern Myanmar, the population density in a sampled area of roughly 3,250 km2 (1,250 sq mi) in a mosaic of tropical broadleaf forest and grassland was estimated to be 0.21–0.44 tigers per 100 km2 (39 sq mi) as of 2009.[75] Population density in mixed deciduous and semi-evergreen forests of Thailand's Huai Kha Khaeng Wildlife Sanctuary was estimated at 2.01 tigers per 100 km2 (39 sq mi); during the 1970s and 1980s, logging and poaching had occurred in the adjacent Mae Wong and Khlong Lan National Parks, where population density was much lower, estimated at only 0.359 tigers per 100 km2 (39 sq mi) as of 2016.[76] Population density in dipterocarp and montane forests in northern Malaysia was estimated at 1.47–2.43 adult tigers per 100 km2 (39 sq mi) in Royal Belum State Park, but 0.3–0.92 adult tigers per 100 km2 (39 sq mi) in the unprotected selectively logged Temengor Forest Reserve.[77]
Behaviour and ecologyTiger bathing
Camera trap data show that tigers in Chitwan National Park avoided locations frequented by people and were more active at night than during day.[78] In Sundarbans National Park, six radio-collared tigers were most active from dawn to early morning and reached their zenith around 7:00 o'clock in the morning.[79] A three-year-long camera trap survey in Shuklaphanta National Park revealed that tigers were most active from dusk until midnight.[80] In northeastern China, tigers were crepuscular and active at night with activity peaking at dawn and dusk; they were largely active at the same time as their prey.[81]
The tiger is a powerful swimmer and easily transverses rivers as wide as 8 km (5.0 mi); it immerses in water, particularly on hot days.[57] In general, it is less capable of climbing trees than many other cats due to its size, but cubs under 16 months old may routinely do so.[82] An adult was recorded climbing 10 m (33 ft) up a smooth pipal tree.[47]
Social spacing
Adult tigers lead largely solitary lives within home ranges or territories, the size of which mainly depends on prey abundance, geographic area and sex of the individual. Males and females defend their home ranges from those of the same sex and the home range of a male encompasses that of multiple females.[47][57] Two females in the Sundarbans had home ranges of 10.6 and 14.1 km2 (4.1 and 5.4 sq mi).[83] In Panna Tiger Reserve, the home ranges of five reintroduced females varied from 53–67 km2 (20–26 sq mi) in winter to 55–60 km2 (21–23 sq mi) in summer and to 46–94 km2 (18–36 sq mi) during the monsoon; three males had 84–147 km2 (32–57 sq mi) large home ranges in winter, 82–98 km2 (32–38 sq mi) in summer and 81–118 km2 (31–46 sq mi) during monsoon seasons.[84] In Sikhote-Alin Biosphere Reserve, 14 females had home ranges 248–520 km2 (96–201 sq mi) and five resident males of 847–1,923 km2 (327–742 sq mi) that overlapped with those of up to five females.[85] When tigresses in the same reserve had cubs of up to four months of age, they reduced their home ranges to stay near their young and steadily enlarged them until their offspring were 13–18 months old.[86]
Bengal tigers spraying urine (above) and rubbing against a tree to mark territory
The tiger is a long-ranging species and individuals disperse over distances of up to 650 km (400 mi) to reach tiger populations in other areas.[87] Young tigresses establish their first home ranges close to their mothers' while males migrate further than their female counterparts.[88] Four radio-collared females in Chitwan dispersed between 0 and 43.2 km (0.0 and 26.8 mi) and 10 males between 9.5 and 65.7 km (5.9 and 40.8 mi).[89] A subadult male lives as a transient in another male's home range until he is older and strong enough to challenge the resident male.[88][90] Tigers mark their home ranges by spraying urine on vegetation and rocks, clawing or scent rubbing trees and marking trails with faeces, anal gland secretions and ground scrapings.[57][91][92][93] Scent markings also allow an individual to pick up information on another's identity. Unclaimed home ranges, particularly those that belonged to a deceased individual, can be taken over in days or weeks.[57]
Male tigers are generally less tolerant of other males within their home ranges than females are of other females. Disputes are usually solved by intimidation rather than fighting. Once dominance has been established, a male may tolerate a subordinate within his range, as long as they do not come near him. The most serious disputes tend to occur between two males competing for a female in oestrus.[94] Though tigers mostly live alone, relationships between individuals can be complex. Tigers are particularly social at kills and a male tiger will sometimes share a carcass with the females and cubs within this home range and unlike male lions, will allow them to feed on the kill before he is finished with it. However, a female is more tense when encountering another female at a kill.[95][96]
CommunicationSiberian tiger baring teeth as a sign of aggressionCaptive Sumatran tiger roaring
During friendly encounters and bonding, tigers rub against each other's bodies.[97] Facial expressions include the "defence threat", which involves a wrinkled face, bared teeth, pulled-back ears and widened pupils.[98][47] Both males and females show a flehmen response, a characteristic curled-lip grimace, when smelling urine markings. Males also use the flehmen to detect the markings made by tigresses in oestrus.[47] Tigers will move their ears around to display the white spots, particularly during aggressive encounters and between mothers and cubs.[99] They also use their tails to signal their mood. To show cordiality, the tail sticks up and sways slowly, while an apprehensive tiger lowers its tail or wags it side-to-side. When calm, the tail hangs low.[100]
Tigers are normally silent but can produce numerous vocalisations.[101][102] They roar to signal their presence to other individuals over long distances. This vocalisation is forced through an open mouth as it closes and can be heard 3 km (1.9 mi) away. They roar multiple times in a row and others respond in kind. Tigers also roar during mating and a mother will roar to call her cubs to her. When tense, tigers moan, a sound similar to a roar but softer and made when the mouth is at least partially closed. Moaning can be heard 400 m (1,300 ft) away.[47][103] Aggressive encounters involve growling, snarling and hissing.[104] An explosive "coughing roar" or "coughing snarl" is emitted through an open mouth and exposed teeth.[47][104][99] In friendlier situations, tigers prusten, a soft, low-frequency snorting sound similar to purring in smaller cats.[105] Tiger mothers communicate with their cubs by grunting, while cubs call back with miaows.[106] When startled, they "woof". They produce a deer-like "pok" sound for unknown reasons, but most often at kills.[107][108]
Hunting and dietBengal tiger attacking a sambar deer in Ranthambore Tiger Reserve
The tiger is a carnivore and an apex predator feeding mainly on large and medium-sized ungulates, with a preference for sambar deer, Manchurian wapiti, barasingha, gaur and wild boar.[109][110][111] Abundance and body weight of prey species are assumed to be the main criteria for the tiger's prey selection, both inside and outside protected areas.[112] It also preys opportunistically on smaller species like monkeys, peafowl and other ground-based birds, porcupines and fish.[47][57] Occasional attacks on Asian elephants and Indian rhinoceroses have also been reported.[113] More often, tigers take the more vulnerable calves.[114] They sometimes prey on livestock and dogs in close proximity to settlements.[47] Tigers occasionally consume vegetation, fruit and minerals for dietary fibre and supplements.[115]
Tigers learn to hunt from their mothers, though the ability to hunt may be partially inborn.[116] Depending on the size of the prey, they typically kill weekly though mothers must kill more often.[54] Families hunt together when cubs are old enough.[117] They search for prey using vision and hearing.[118] A tiger will also wait at a watering hole for prey to come by, particularly during hot summer days.[119][120] It is an ambush predator and when approaching potential prey, it crouches with the head lowered and hides in foliage. It switches between creeping forward and staying still. A tiger may even doze off and can stay in the same spot for as long as a day, waiting for prey and launch an attack when the prey is close enough,[121] usually within 30 m (98 ft).[54] If the prey spots it before then, the cat does not pursue further.[119] A tiger can sprint 56 km/h (35 mph) and leap 10 m (33 ft);[122][123] it is not a long-distance runner and gives up a chase if prey outpaces it over a certain distance.[119]
Two Bengal tigers attacking a wild boar in Kanha Tiger Reserve
The tiger attacks from behind or at the sides and tries to knock the target off balance. It latches onto prey with its forelimbs, twisting and turning during the struggle and tries to pull it to the ground. The tiger generally applies a bite to the throat until its victim dies of strangulation.[47][124][125][126] It has an average bite force at the canine tips of 1234.3 newtons.[127] Holding onto the throat puts the cat out of reach of horns, antlers, tusks and hooves.[124][128] Tigers are adaptable killers and may use other methods, including ripping the throat or breaking the neck. Large prey may be disabled by a bite to the back of the hock, severing the tendon. Swipes from the large paws are capable of stunning or breaking the skull of a water buffalo.[129] They kill small prey with a bite to the back of the neck or head.[130][54] Estimates of the success rate for hunting tigers range from a low of 5% to a high of 50%. They are sometimes killed or injured by large or dangerous prey like gaur, buffalo and boar.[54]
Tigers typically move kills to a private, usually vegetated spot no further than 183 m (600 ft), though they have been recorded dragging them 549 m (1,801 ft). They are strong enough to drag the carcass of a fully grown buffalo for some distance. They rest for a while before eating and can consume as much as 50 kg (110 lb) of meat in one session, but feed on a carcass for several days, leaving little for scavengers.[131]
CompetitorsAn 1807 illustration of dholes attacking a tiger
In much of their range, tigers share habitat with leopards and dholes. They typically dominate both of them, though with dholes it depends on their pack size. Interactions between the three predators involve chasing, stealing kills and direct killing.[132] Large dhole packs may kill tigers.[133] Tigers, leopards and dholes coexist by hunting different sized prey.[134] In Nagarhole National Park, the average weight for tiger kills was found to be 91.5 kg (202 lb), compared to 37.6 kg (83 lb) for leopards and 43.4 kg (96 lb) for dholes.[135] In Kui Buri National Park, following a reduction in prey numbers, tigers continued to kill favoured prey while leopards and dholes increased their consumption of small prey.[110]
Both leopards and dholes can live successfully in tiger habitat when there is abundant food and vegetation cover.[134][136] Otherwise, they appear to be less common where tigers are numerous. The recovery of the tiger population in Rajaji National Park during the 2000s led to a reduction in leopard population densities.[137] Similarly, at two sites in central India the size of dhole packs was negatively correlated with tiger densities.[138] Leopard and dhole distribution in Kui Buri correlated with both prey access and tiger scarcity.[139] In Jigme Dorji National Park, tigers were found to inhabit the deeper parts of forests while the smaller predators were pushed closer to the fringes.[140]
Reproduction and life cycle"Tiger cub" redirects here. For other uses, see Tiger Cub.A Bengal tiger family in Kanha Tiger Reserve
The tiger generally mates all year round, particularly between November and April. A tigress is in oestrus for three to six days at a time, separated by three to nine week intervals.[47] A resident male mates with all the females within his home range, who signal their receptiveness by roaring and marking.[141][142] Younger, transient males are also attracted, leading to a fight in which the more dominant, resident male drives the usurper off.[143][141] During courtship, the male is cautious with the female as he waits for her to show signs she is ready to mate. She signals to him by positioning herself in lordosis with her tail to the side. Copulation typically lasts no more than 20 seconds, with the male biting the female by the scruff of her neck. After it is finished, the male quickly pulls away as the female may turn and slap him.[141] Tiger pairs may stay together for up to four days and mate multiple times.[144] Gestation lasts around or over three months.[47]
A tigress gives birth in a secluded location, be it in dense vegetation, in a cave or under a rocky shelter.[145] Litters consist of as many as seven cubs, but two or three are more typical.[143][145] Newborn cubs weigh 785–1,610 g (27.7–56.8 oz) and are blind and altricial.[145] The mother licks and cleans her cubs, suckles them and viciously defends them from any potential threat.[143] Cubs open their eyes at the age of three to 14 days and their vision becomes clear after a few more weeks.[145] They can leave the denning site after two months and around the same time they start eating meat.[143][146] The mother only leaves them alone to hunt and even then she does not travel far.[147] When she suspects an area is no longer safe, she moves her cubs to a new spot, transporting them one by one by grabbing them by the scruff of the neck with her mouth.[148] A tigress in Sikhote-Alin Biosphere Reserve maximised the time spent with her cubs by reducing her home range, killing larger prey and returning to her den more rapidly than without cubs; when the cubs started to eat meat, she took them to kill sites, thereby optimising their protection and access to food.[149] In the same reserve, one of 21 cubs died in over eight years of monitoring and mortality did not differ between male and female juveniles.[150] Tiger monitoring over six years in Ranthambore Tiger Reserve indicated an average annual survival rate of around 85 percent for 74 male and female cubs; survival rate increased to 97 percent for both males and female juveniles of one to two years of age.[151] Causes of cub mortality include predators, floods, fires, death of the mother and fatal injuries.[148][152][153][151]
A Siberian tigress with her cub at Buffalo Zoo
After around two months, the cubs are able to follow their mother. They still hide in vegetation when she goes hunting. Young bond through play fighting and practice stalking. A hierarchy develops in the litter, with the biggest cub, often a male, being the most dominant and the first to eat its fill at a kill.[154] Around the age of six months, cubs are fully weaned and have more freedom to explore their environment. Between eight and ten months, they accompany their mother on hunts.[117] A cub can make a kill as early as 11 months and reach independence as a juvenile of 18 to 24 months of age; males become independent earlier than females.[155][151] Radio-collared tigers in Chitwan started leaving their natal areas at the age of 19 months.[89] Young females are sexually mature at three to four years, whereas males are at four to five years.[47] Generation length of the tiger is about 7–10 years.[156] Wild Bengal tigers live 12–15 years.[157] Data from the International Tiger Studbook 1938–2018 indicate that captive tigers lived up to 19 years.[158]
The father does not play a role in raising the young, but he encounters and interacts with them. The resident male appears to visit the female–cub families within his home range. They socialise and even share kills.[159][160] One male was recorded looking after cubs whose mother had died.[161] By defending his home range, the male protects the females and cubs from other males.[162] When a new male takes over, dependent cubs are at risk of infanticide as the male attempts to sire his own young with the females.[163] A seven-year long study in Chitwan National Park revealed that 12 of 56 detected cubs and juveniles were killed by new males taking over home ranges.[153]
Health and diseases
Tigers are recorded as hosts for various parasites including tapeworms like Diphyllobothrium erinacei, Taenia pisiformis in India and nematodes like Toxocara species in India and Physaloptera preputialis, Dirofilaria ursi and Uiteinarta species in Siberia.[47] Canine distemper is known to occur in Siberian tigers.[164] A morbillivirus infection was the likely cause of death of a tigress in the Russian Far East that was also tested positive for feline panleukopenia and feline coronavirus.[165] Blood samples from 11 adult tigers in Nepal showed antibodies for canine parvovirus-2, feline herpesvirus, feline coronavirus, leptospirosis and Toxoplasma gondii.[166]
Threats
The tiger has been listed as Endangered on the IUCN Red List since 1986 and the global tiger population is thought to have continuously declined from an estimated population of 5,000–8,262 tigers in the late 1990s to 3,726–5,578 individuals estimated as of 2022.[1] During 2001–2020, landscapes where tigers live declined from 1,025,488 km2 (395,943 sq mi) to 911,901 km2 (352,087 sq mi).[65] Habitat destruction, habitat fragmentation and poaching for fur and body parts are the major threats that contributed to the decrease of tiger populations in all range countries.[1]
Protected areas in central India are highly fragmented due to linear infrastructure like roads, railway lines, transmission lines, irrigation channels and mining activities in their vicinity.[167] In the Tanintharyi Region of southern Myanmar, deforestation coupled with mining activities and high hunting pressure threatens the tiger population.[168] In Thailand, nine of 15 protected areas hosting tigers are isolated and fragmented, offering a low probability for dispersal between them; four of these have not harboured tigers since about 2013.[169] In Peninsular Malaysia, 8,315.7 km2 (3,210.7 sq mi) of tiger habitat was cleared during 1988–2012, most of it for industrial plantations.[170] Large-scale land acquisitions of about 23,000 km2 (8,900 sq mi) for commercial agriculture and timber extraction in Cambodia contributed to the fragmentation of potential tiger habitat, especially in the Eastern Plains.[171] Inbreeding depression coupled with habitat destruction, insufficient prey resources and poaching is a threat to the small and isolated tiger population in the Changbai Mountains along the China–Russia border.[172] In China, tigers became the target of large-scale 'anti-pest' campaigns in the early 1950s, where suitable habitats were fragmented following deforestation and resettlement of people to rural areas, who hunted tigers and prey species. Though tiger hunting was prohibited in 1977, the population continued to decline and is considered extinct in South China since 2001.[173][174]
A Javan tiger skin, 1915
Tiger populations in India have been targeted by poachers since the 1990s and were extirpated in two tiger reserves in 2005 and 2009.[175] Between March 2017 and January 2020, 630 activities of hunters using snares, drift nets, hunting platforms and hunting dogs were discovered in a reserve forest of about 1,000 km2 (390 sq mi) in southern Myanmar.[176] Nam Et-Phou Louey National Park was considered the last important site for the tiger in Laos, but it has not been recorded there at least since 2013; this population likely fell victim to indiscriminate snaring.[177] Anti-poaching units in Sumatra's Kerinci Seblat landscape removed 362 tiger snare traps and seized 91 tiger skins during 2005–2016; annual poaching rates increased with rising skin prices.[178] Poaching is also the main threat to the tiger population in far eastern Russia, where logging roads facilitate access for poachers and people harvesting forest products that are important for prey species to survive in winter.[179]
Body parts of 207 tigers were detected during 21 surveys in 1991–2014 in two wildlife markets in Myanmar catering to customers in Thailand and China.[180] During the years 2000–2022, at least 3,377 tigers were confiscated in 2,205 seizures in 28 countries; seizures encompassed 665 live and 654 dead individuals, 1,313 whole tiger skins, 16,214 body parts like bones, teeth, paws, claws, whiskers and 1.1 t (1.1 long tons; 1.2 short tons) of meat; 759 seizures in India encompassed body parts of 893 tigers; and 403 seizures in Thailand involved mostly captive-bred tigers.[181] Seizures in Nepal between January 2011 and December 2015 obtained 585 pieces of tiger body parts and two whole carcasses in 19 districts.[182] Seizure data from India during 2001–2021 indicate that tiger skins were the most often traded body parts, followed by claws, bones and teeth; trafficking routes mainly passed through the states of Maharashtra, Karnataka, Tamil Nadu and Assam.[183] A total of 292 illegal tiger parts were confiscated at US ports of entry from personal baggage, air cargo and mail between 2003 and 2012.[184]
Demand for tiger parts for use in traditional Chinese medicine has also been cited as a major threat to tiger populations.[185] Interviews with local people in the Bangladeshi Sundarbans revealed that they kill tigers for local consumption and trade of skins, bones and meat, in retaliation for attacks by tigers and for excitement.[186] Tiger body parts like skins, bones, teeth and hair are consumed locally by wealthy Bangladeshis and are illegally trafficked from Bangladesh to 15 countries including India, China, Malaysia, Korea, Vietnam, Cambodia, Japan and the United Kingdom via land borders, airports and seaports.[187] Tiger bone glue is the prevailing tiger product purchased for medicinal purposes in Hanoi and Ho Chi Minh City.[188] "Tiger farm" facilities in China and Southeast Asia breed tigers for their parts, but these appear to make the threat to wild populations worse by increasing the demand for tiger products.[189]
Local people killing tigers in retaliation for attacking and preying on livestock is a threat in several tiger range countries, as this consequence of human–wildlife conflict also contributes to the decline of the population.[190][191][192][193][194]
ConservationMain article: Tiger conservationFurther information: 21st Century TigerGlobal wild tiger populationCountryYearEstimate India20223,167–3,682[195] Russia2022573–600[196] Indonesia2022393[196] Nepal2022316–355[197] Thailand2022148–189[196] Malaysia2022<150[196] Bhutan2022131[198] Bangladesh2022118–122[199] China2022>60[196] Myanmar202228[196]Total5,638–5,899
Internationally, the tiger is protected under CITES Appendix I, banning trade of live tigers and their body parts.[1] In Russia, hunting the tiger has been banned since 1952.[200] In Bhutan, it has been protected since 1969 and enlisted as totally protected since 1995.[201] Since 1972, it has been afforded the highest protection level under India's Wild Life (Protection) Act, 1972.[202] In Nepal and Bangladesh, it has been protected since 1973.[202][187] Since 1976, it has been totally protected under Malaysia's Protection of Wild Life Act,[203] and the country's Wildlife Conservation Act enacted in 2010 increased punishments for wildlife-related crimes.[196] In Indonesia, it has been protected since 1990.[204] In China, the trade in tiger body parts was banned in 1993.[205] The Thai Wildlife Preservation and Protection Act was enacted in 2019 to combat poaching and trading of body parts.[206]
In 1973, the National Tiger Conservation Authority and Project Tiger were founded in India to gain public support for tiger conservation.[175] Since then, 53 tiger reserves covering an area of 75,796 km2 (29,265 sq mi) have been established in the country up to 2022.[195] Myanmar's national tiger conservation strategy developed in 2003 comprises management tasks such as restoration of degraded habitats, increasing the extent of protected areas and wildlife corridors, protecting tiger prey species, thwarting tiger killing and illegal trade of its body parts and promoting public awareness through wildlife education programmes.[207] Bhutan's first Tiger Action Plan implemented during 2006–2015 revolved around habitat conservation, human–wildlife conflict management, education and awareness; the second Action Plan aimed at increasing the country's tiger population by 20% until 2023 compared to 2015.[201] In 2009, the Bangladesh Tiger Action Plan was initiated to stabilise the country's tiger population, maintain habitat and a sufficient prey base, improve law enforcement and foster cooperation between governmental agencies responsible for tiger conservation.[208] The Thailand Tiger Action Plan ratified in 2010 envisioned increasing the country's tiger populations by 50% in the Western Forest Complex and Dong Phayayen–Khao Yai Forest Complex and reestablish populations in three potential landscapes until 2022.[209] The Indonesian National Tiger Recovery Program ratified in 2010 aimed at increasing the Sumatran tiger population by 2022.[210] The third strategic and action plan for the conservation of the Sumatran tiger for the years 2020–2030 revolves around strengthening management of small tiger population units of less than 20 mature individuals and connectivity between 13 forest patches in North Sumatra and West Sumatra provinces.[211]
Wild Sumatran tiger caught by camera trap
Increases in anti-poaching patrol efforts in four Russian protected areas during 2011–2014 contributed to reducing poaching, stabilising the tiger population and improving protection of ungulate populations.[212] Poaching and trafficking were declared to be moderate and serious crimes in 2019.[196] Anti-poaching operations were also established in Nepal in 2010, with increased cooperation and intelligence sharing between agencies. These policies have led to many years of "zero poaching" and the country's tiger population has doubled in a decade.[196] Anti-poaching patrols in the 1,200 km2 (460 sq mi) large core area of Taman Negara lead to a decrease of poaching frequency from 34 detected incidents in 2015–2016 to 20 incidents during 2018–2019; the arrest of seven poaching teams and removal of snares facilitated the survival of three resident female tigers and at least 11 cubs.[213] Army and police officers are deployed for patrolling together with staff of protected areas in Malaysia.[196]
Wildlife corridors are important conservation measures as they facilitate tiger populations to connect between protected areas; tigers use at least nine corridors that were established in the Terai Arc Landscape and Sivalik Hills in both Nepal and India.[214] Corridors in forested areas with low human encroachment are highly suitable.[215][216] In West Sumatra, 12 wildlife corridors were identified as high priority for mitigating human–wildlife conflicts.[217] In 2019, China and Russia signed a memorandum of understanding for transboundary cooperation between two protected areas, Northeast China Tiger and Leopard National Park and Land of the Leopard National Park, that includes the creation of wildlife corridors and bilateral monitoring and patrolling along the Sino-Russian border.[218]
Rescued and rehabilitated problem tigers and orphaned tiger cubs have been released into the wild and monitored in India, Sumatra and Russia.[84][219][220] In Kazakhstan, habitat restoration and reintroduction of prey species in Ile-Balkash Nature Reserve have progressed and tiger reintroduction is planned for 2025.[221] Reintroduction of tigers is considered possible in eastern Cambodia, once management of protected areas is improved and forest loss stabilized.[222] South China tigers are kept and bred in Chinese zoos, with plans to reintroduce their offspring into remote protected areas.[15][116] Coordinated breeding programs among zoos have led to enough genetic diversity in tigers to act as "insurance against extinction in the wild".[223]
Relationship with humansTiger hunting on elephant-back in India, 1808HuntingMain article: Tiger hunting
Tigers have been hunted by humans for millennia, as indicated by a painting on the Bhimbetka rock shelters in India that is dated to 5,000–6,000 years ago. They were hunted throughout their range in Asia, chased on horseback, elephant-back or even with sled dogs and killed with spears and later firearms. Such hunts were conducted both by native governments and empires like the Mughal Empire, as well as European colonists. Tigers were often hunted as trophies and because of their perceived danger.[224] An estimated 80,000 tigers were killed between 1875 and 1925.[225][226]
AttacksMain article: Tiger attackA Bengal tiger in the Sundarbans
In most areas, tigers avoid humans, but attacks are a risk wherever people coexist with them.[227][228] Dangerous encounters are more likely to occur in edge habitats between wild and agricultural areas.[227] Most attacks on humans are defensive, including protection of young; however, tigers do sometimes see people as prey.[228] Man-eating tigers tend to be old and disabled.[57] Tigers driven from their home ranges are also at risk of turning to man-eating.[229]
At the beginning of the 20th century, the Champawat Tiger was responsible for over 430 human deaths in Nepal and India before she was shot by Jim Corbett.[230] This tigress suffered from broken teeth and was unable to kill normal prey. Modern authors speculate that sustaining on meagre human flesh forced the cat to kill more and more.[231] Tiger attacks were particularly high in Singapore during the mid-19th century, when plantations expanded into the tiger's habitat.[232] In the 1840s, the number of deaths in the area ranged from 200 to 300 annually.[233] Tiger attacks in the Sundarbans caused 1,396 human deaths in the period 1935–2006 according to official records of the Bangladesh Forest Department.[234] Victims of these attacks are local villagers who enter the tiger's domain to collect resources like wood and honey. Fishermen have been particularly common targets. Methods to counter tiger attacks have included face masks worn backwards, protective clothes, sticks and carefully stationed electric dummies.[235]
CaptivityTiger at Big Cat Rescue in 2014Publicity photo of animal trainer Gunther Gebel-Williams with several of his trained tigers, c. 1969
Tigers have been kept in captivity since ancient times. In ancient Rome, tigers were displayed in amphitheatres; they were slaughtered in venatio hunts and used to kill criminals.[236] The Mongol ruler Kublai Khan is reported to have kept tigers in the 13th century. Starting in the Middle Ages, tigers were being kept in European menageries.[237] Tigers and other exotic animals were mainly used for the entertainment of elites but from the 19th century onward, they were exhibited more to the public. Tigers were particularly big attractions and their captive population soared.[238] In 2020, there were over 8,000 captive tigers in Asia, over 5,000 in the US and no less than 850 in Europe.[239] There are more tigers in captivity than in the wild.[223] Captive tigers may display stereotypical behaviours such as pacing or inactivity. Modern zoos are able to reduce such behaviours with exhibits designed so the animals can move between separate but connected enclosures.[240] Enrichment items are also important for the cat's welfare and the stimulation of its natural behaviours.[241]
Tigers have played prominent roles in circuses and other live performances. Ringling Bros included many tiger tamers in the 20th century including Mabel Stark, who became a big draw and had a long career. She was well known for being able to control the tigers despite being a small woman; using "manly" tools like whips and guns. Another trainer was Clyde Beatty, who used chairs, whips and guns to provoke tigers and other beasts into acting fierce and allowed him to appear courageous. He would perform with as many as 40 tigers and lions in one act. From the 1960s onward, trainers like Gunther Gebel-Williams would use gentler methods to control their animals. Sara Houcke was dubbed "the Tiger Whisperer" as she trained the cats to obey her by whispering to them.[242] Siegfried & Roy became famous for performing with white tigers in Las Vegas. The act ended in 2003 when a tiger attacked Roy during a performance.[243] In 2009, tigers were the most traded circus animals.[244] The use of tigers and other animals in shows eventually declined in many countries due to pressure from animal rights groups and greater desires from the public to see them in more natural settings. Several countries restrict or ban such acts.[245]
Tigers have become popular in the exotic pet trade, particularly in the United States[246] where only 6% of the captive tiger population in 2020 were being housed in zoos and other facilities approved by the Association of Zoos and Aquariums.[189] Private collectors are thought to be ill-equipped to provide proper care for tigers, which compromises their welfare. They can also threaten public safety by allowing people to interact with them.[189] The keeping of tigers and other big cats by private people was banned in the US in 2022.[247] Most countries in the European Union have banned breeding and keeping tigers outside of licensed zoos and rescue centres, but some still allow private holdings.
The lion (Panthera leo) is a large cat of the genus Panthera, native to Africa and India. It has a muscular, broad-chested body; a short, rounded head; round ears; and a dark, hairy tuft at the tip of its tail. It is sexually dimorphic; adult male lions are larger than females and have a prominent mane. It is a social species, forming groups called prides. A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on medium-sized and large ungulates. The lion is an apex and keystone predator.
The lion inhabits grasslands, savannahs, and shrublands. It is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight. During the Neolithic period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India. It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have declined by about 43% since the early 1990s. Lion populations are untenable outside designated protected areas. Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes for concern.
One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in literature and films. Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoological gardens across the world since the late 18th century. Cultural depictions of lions were prominent in Ancient Egypt, and depictions have occurred in virtually all ancient and medieval cultures in the lion's historic and current range.
Etymology
The English word lion is derived via Anglo-Norman liun from Latin leōnem (nominative: leō), which in turn was a borrowing from Ancient Greek λέων léōn. The Hebrew word לָבִיא lavi may also be related.[4] The generic name Panthera is traceable to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'.[5]
TaxonomyThe upper cladogram is based on the 2006 study,[6][7] the lower one on the 2010[8] and 2011[9] studies.
Felis leo was the scientific name used by Carl Linnaeus in 1758, who described the lion in his work Systema Naturae.[3] The genus name Panthera was coined by Lorenz Oken in 1816.[10] Between the mid-18th and mid-20th centuries, 26 lion specimens were described and proposed as subspecies, of which 11 were recognised as valid in 2005.[1] They were distinguished mostly by the size and colour of their manes and skins.[11]
SubspeciesRange map showing distribution of subspecies and clades
In the 19th and 20th centuries, several lion type specimens were described and proposed as subspecies, with about a dozen recognised as valid taxa until 2017.[1] Between 2008 and 2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion populations, and P. l. persica for the Asiatic lion population.[2][12][13] In 2017, the Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy, and recognises two subspecies based on results of several phylogeographic studies on lion evolution, namely:[14]
P. l. leo (Linnaeus, 1758) − the nominate lion subspecies includes the Asiatic lion, the regionally extinct Barbary lion, and lion populations in West and northern parts of Central Africa.[14] Synonyms include P. l. persica (Meyer, 1826), P. l. senegalensis (Meyer, 1826), P. l. kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).[1] Multiple authors referred to it as 'northern lion' and 'northern subspecies'.[15][16]P. l. melanochaita (Smith, 1842) − includes the extinct Cape lion and lion populations in East and Southern African regions.[14] Synonyms include P. l. somaliensis (Noack 1891), P. l. massaica (Neumann, 1900), P. l. sabakiensis (Lönnberg, 1910), P. l. bleyenberghi (Lönnberg, 1914), P. l. roosevelti (Heller, 1914), P. l. nyanzae (Heller, 1914), P. l. hollisteri (Allen, 1924), P. l. krugeri (Roberts, 1929), P. l. vernayi (Roberts, 1948), and P. l. webbiensis (Zukowsky, 1964).[1][11] It has been referred to as 'southern subspecies' and 'southern lion'.[16]
However, there seems to be some degree of overlap between both groups in northern Central Africa. DNA analysis from a more recent study indicates that Central African lions are derived from both northern and southern lions, as they cluster with P. leo leo in mtDNA-based phylogenies whereas their genomic DNA indicates a closer relationship with P. leo melanochaita.[17]
Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East Africa. Researchers, therefore, assume Ethiopia is a contact zone between the two subspecies.[18] Genome-wide data of a wild-born historical lion sample from Sudan showed that it clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. l. melanochaita. This result suggested that the taxonomic position of lions in Central Africa may require revision.[19]
Fossil recordsSkull of an American lion on display at the National Museum of Natural History
Other lion subspecies or sister species to the modern lion existed in prehistoric times:[20]
P. l. sinhaleyus was a fossil carnassial excavated in Sri Lanka, which was attributed to a lion. It is thought to have become extinct around 39,000 years ago.[21]P. fossilis was larger than the modern lion and lived in the Middle Pleistocene. Bone fragments were excavated in caves in the United Kingdom, Germany, Italy and Czech Republic.[22][23]P. spelaea, or the cave lion, lived in Eurasia and Beringia during the Late Pleistocene. It became extinct due to climate warming or human expansion latest by 11,900 years ago.[24] Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska.[25] It likely derived from P. fossilis,[26] and was genetically isolated and highly distinct from the modern lion in Africa and Eurasia.[27][26] It is depicted in Paleolithic cave paintings, ivory carvings, and clay busts.[28]P. atrox, or the American lion, ranged in the Americas from Canada to possibly Patagonia during the Late Pleistocene.[29] It diverged from the cave lion around 165,000 years ago.[30] A fossil from Edmonton dates to 11,355 ± 55 years ago.[31]Evolutionred Panthera spelaea
blue Panthera atrox
green Panthera leo
Maximal range of the modern lion
and its prehistoric relatives
in the late Pleistocene
The Panthera lineage is estimated to have genetically diverged from the common ancestor of the Felidae around 9.32 to 4.47 million years ago to 11.75 to 0.97 million years ago.[6][32][33] Results of analyses differ in the phylogenetic relationship of the lion; it was thought to form a sister group with the jaguar that diverged 3.46 to 1.22 million years ago,[6] but also with the leopard that diverged 3.1 to 1.95 million years ago[8][9] to 4.32 to 0.02 million years ago. Hybridisation between lion and snow leopard ancestors possibly continued until about 2.1 million years ago.[33] The lion-leopard clade was distributed in the Asian and African Palearctic since at least the early Pliocene.[34] The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old.[32]
Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on mutation rate per generation time of the modern lion. There is no evidence for gene flow between the two lineages, indicating that they did not share the same geographic area.[19] The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents.[27][35][36] The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago.[37] They shared a common ancestor probably between 98,000 and 52,000 years ago.[19] Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia between 38,800 and 8,300 years ago.[37]
Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa. Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.[38] A whole genome-wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This phenomenon indicates that Central Africa was a melting pot of lion populations after they had become isolated, possibly migrating through corridors in the Nile Basin during the early Holocene.[19]
HybridsFurther information: Panthera hybrid
In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose.[39][40] The liger is bigger than a lion and a tiger, whereas most tigons are relatively small compared to their parents because of reciprocal gene effects.[41][42] The leopon is a hybrid between a lion and leopard.[43]
DescriptionA tuft at the end of the tail is a distinct characteristic of the lion.Skeleton
The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck, and round ears; males have broader heads. The fur varies in colour from light buff to silvery grey, yellowish red, and dark brown. The colours of the underparts are generally lighter. A new-born lion has dark spots, which fade as the cub reaches adulthood, although faint spots may still be seen on the legs and underparts.[44][45] The tail of all lions ends in a dark, hairy tuft that, in some lions, conceals an approximately 5 mm (0.20 in)-long, hard "spine" or "spur" composed of dermal papillae.[46] The functions of the spur are unknown. The tuft is absent at birth and develops at around 5+1⁄2 months of age. It is readily identifiable at the age of seven months.[47]
Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened and has a slightly shorter postorbital region and broader nasal openings than those of the tiger. Due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species.[48][49]
The skeletal muscles of the lion make up 58.8% of its body weight and represent the highest percentage of muscles among mammals.[50][51] The lion has a high concentration of fast twitch muscle fibres, giving them quick bursts of speed but less stamina.[52][53]
The male lion's mane is the most recognisable feature of the species.[11] It may have evolved around 320,000–190,000 years ago.[62] It grows downwards and backwards, covering most of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow, rust, and black hairs.[45] Mutations in the genes microphthalmia-associated transcription factor and tyrosinase are possibly responsible for the colour of manes.[63][64] It starts growing when lions enter adolescence, when testosterone levels increase, and reach their full size at around four years old.[65] Cool ambient temperatures in European and North American zoos may result in a heavier mane.[66] On average, Asiatic lions have sparser manes than African lions.[67]
This feature likely evolved to signal the fitness of males to females. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress.[68][69] The core body temperature does apparently not increase regardless of sex, season, feeding time, length and colour of mane, but only surface temperature is affected.[70] Unlike in other felid species, female lions consistently interact with multiple males at once.[71] Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.[72][73] During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters. Injured lions also begin to lose their manes.[74]
Almost all male lions in Pendjari National Park are either maneless or have very short manes.[75] Maneless lions have also been reported in Senegal, in Sudan's Dinder National Park and in Tsavo East National Park, Kenya.[76] Castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production.[77] Rarely, both wild and captive lionesses have manes.[78][79] Increased testosterone may be the cause of maned lionesses reported in northern Botswana.[80]
Colour variationFurther information: White lion
The white lion is a rare morph with a genetic condition called leucism, which is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool. Nevertheless, 17 births have been recorded in five prides between 2007 and 2015.[81] White lions are selected for breeding in captivity.[82] They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts.[83]
Distribution and habitatLion in Gir National Park, India
African lions live in scattered populations across sub-Saharan Africa. The lion prefers grassy plains and savannahs, scrub bordering rivers, and open woodlands with bushes. It rarely enters closed forests. On Mount Elgon, the lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.[44] Savannahs with an annual rainfall of 300 to 1,500 mm (12 to 59 in) make up the majority of lion habitat in Africa, estimated at 3,390,821 km2 (1,309,203 sq mi) at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa.[84] The Asiatic lion now survives only in and around Gir National Park in Gujarat, western India. Its habitat is a mixture of dry savannah forest and very dry, deciduous scrub forest.[12]
Historical range
In Africa, the range of the lion originally spanned most of the central African rainforest zone and the Sahara desert.[85] In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.[86][84][87]
During the mid-Holocene, around 8,000-6,000 years ago, the range of lions expanded into Southeastern and Eastern Europe, partially re-occupying the range of the now extinct cave lion.[88] In Hungary, the modern lion was present from about 4,500 to 3,200 years Before Present.[89] In Ukraine, the modern lion was present from about 6,400 to 2,000 years Before Present.[88] In Greece, it was common, as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100.[44]
In Asia the lion once ranged in regions where climatic conditions supported an abundance of prey.[90] It was present in the Caucasus until the 10th century.[49] It lived in Palestine until the Middle Ages and in Southwest Asia until the late 19th century. By the late 19th century, it had been extirpated in most of Turkey.[91] The last live lion in Iran was sighted in 1942, about 65 km (40 mi) northwest of Dezful,[92] although the corpse of a lioness was found on the banks of the Karun river in Khuzestan province in 1944.[93] It once ranged from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India.[94]
Behaviour and ecology
Lions spend much of their time resting; they are inactive for about twenty hours per day.[95] Although lions can be active at any time, their activity generally peaks after dusk with a period of socialising, grooming, and defecating. Intermittent bursts of activity continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and fifty minutes eating.[96]
Group organisationLion pride in Etosha National ParkA lioness (left) and two males in Masai Mara
The lion is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a "pride". Groups of male lions are called "coalitions".[97] Females form the stable social unit in a pride and do not tolerate outside females.[98] The majority of females remain in their birth prides while all males and some females will disperse.[99] The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have been observed.[100] The sole exception to this pattern is the Tsavo lion pride that always has just one adult male.[101] Prides act as fission–fusion societies, and members will split into subgroups that keep in contact with roars.[102]
Nomadic lions range widely and move around sporadically, either in pairs or alone.[97] Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.[103] Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them.[104] Males spend years in a nomadic phase before gaining residence in a pride.[105] A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[106] In Kruger National Park, dispersing male lions move more than 25 km (16 mi) away from their natal pride in search of their own territory. Female lions stay closer to their natal pride. Therefore, female lions in an area are more closely related to each other than male lions in the same area.[107]
The evolution of sociability in lions was likely driven both by high population density and the clumped resources of savannah habitats. The larger the pride, the more high-quality territory they can defend; "hotspots" being near river confluences, where the cats have better access to water, prey and shelter (via vegetation).[108][109] The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".[97] Males associated with a pride patrol the fringes.[45] Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind.[110] Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.[111] Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses.[112] The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.[103] Dominance hierarchies do not appear to exist among individuals of either sex in a pride.[113]
Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming a loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs. Female and male lions associate only when mating.[114] Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently.[115]
Hunting and dietA skeletal mount of a lion attacking a common eland, on display at The Museum of OsteologyFour lionesses catching a buffalo in the SerengetiLions feeding on a zebra
The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.[116][117] Its prey consists mainly of medium-sized to large ungulates, particularly blue wildebeest, plains zebra, African buffalo, gemsbok and giraffe. It also frequently takes common warthog despite it being much smaller.[118] In India, chital and sambar deer are the most common wild prey,[45][118][119] while livestock contributes significantly to lion kills outside protected areas.[120] It usually avoids fully grown adult elephants, rhinoceros and hippopotamus and small prey like dik-dik, hyraxes, hares and monkeys. Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them.[121]
Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two.[122] Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.[103] In Chobe National Park, large prides have been observed hunting African bush elephants up to around 15 years old in exceptional cases, with the victims being calves, juveniles, and even subadults.[123][124] In typical group hunts, each lioness has a favoured position in the group, either stalking prey on the "wing", then attacking, or moving a smaller distance in the centre of the group and capturing prey fleeing from other lionesses. Males attached to prides do not usually participate in group hunting.[125] Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.[126] They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own. Moderately-sized hunting groups generally have higher success rates than lone females and larger groups.[127]
Lions are not particularly known for their stamina. For instance, a lioness's heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight.[128] Thus, lions run quickly only in short bursts at about 48–59 km/h (30–37 mph) and need to be close to their prey before starting the attack.[129] They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[130] One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph).[131] The lion accelerates at the start of the chase by 9.5 m/s², whereas zebras, wildebeest and Thomson's gazelle accelerate by 5 m/s², 5.6 m/s² and 4.5 m/s², respectively; acceleration appears to be more important than steady displacement speed in lion hunts.[132] The lion's attack is short and powerful; it attempts to catch prey with a fast rush and final leap, usually pulls it down by the rump, and kills with a clamping bite to the throat or muzzle. It can hold the prey's throat for up to 13 minutes, until the prey stops moving.[133] It has a bite force from 1593.8 to 1768 Newtons at the canine tip and up 4167.6 Newtons at the carnassial notch.[134][135][136]
Male lion and cub with mostly eaten buffalo carcass in Sabi Sand Game Reserve
Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.[137] They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers.[103] Large kills are shared more widely among pride members.[138] An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).[139] Lions gorge themselves and eat up to 30 kg (66 lb) in one session.[93] If it is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard.[137] Lions defend their kills from scavengers such as vultures and hyenas.[103]
Lions scavenge on carrion when the opportunity arises, scavenging animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep a constant lookout for circling vultures, which indicate the death or distress of an animal.[140] Most carrion on which