Tiger

Tiger
Temporal range: Early Pleistocene – Present
A Bengal tigress in Kanha Tiger Reserve, India
CITES Appendix I (CITES)[1]
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species:
P. tigris
Binomial name
Panthera tigris
(Linnaeus, 1758)[2]
Subspecies
Tiger distribution as of 2022
Synonyms[3]

The tiger (Panthera tigris) is the largest living cat species and a member of the genus Panthera. It is most recognisable for its black stripes on orange fur with a white underside. An apex predator, it primarily preys on ungulates, such as deer and wild boar. It is territorial and generally a solitary but social predator, requiring large contiguous areas of habitat to support its requirements for prey and rearing of its offspring. Tiger cubs stay with their mother for about two years and then become independent, leaving their mother's home range to establish their own.

The tiger was first scientifically described in 1758. It once ranged widely from the Eastern Anatolia Region in the west to the Amur River basin in the east, and in the south from the foothills of the Himalayas to Bali in the Sunda Islands. Since the early 20th century, tiger populations have lost at least 93% of their historic range and have been extirpated from Western and Central Asia, the islands of Java and Bali, and in large areas of Southeast and South Asia and China. What remains of the range where tigers still roam free is fragmented, stretching in spots from Siberian temperate forests to subtropical and tropical forests on the Indian subcontinent, Indochina and a single Indonesian island, Sumatra.

The tiger is listed as Endangered on the IUCN Red List. India hosts the largest tiger population. Major reasons for population decline are habitat destruction, habitat fragmentation and poaching. Tigers are also victims of human–wildlife conflict, due to encroachment in countries with a high human population density.

The tiger is among the most recognisable and popular of the world's charismatic megafauna. It featured prominently in the ancient mythology and folklore of cultures throughout its historic range and continues to be depicted in modern films and literature, appearing on many flags, coats of arms and as mascots for sporting teams. The tiger is the national animal of India, Bangladesh, Malaysia and South Korea.

Etymology

The Old English tigras derive from Old French tigre, from Latin tigris. This was a borrowing of Classical Greek τίγρις 'tigris'.[4][5] The ultimate origin of the word is uncertain.[6] Ancient Greek geographer Strabo suggested an Armenian origin.[7] One popular idea, believed in the 16th and 17th centuries, is tiger was a transliteration of the Middle Persian tigr, meaning 'arrow', from which the name of the river Tigris may also have been derived. Thus, the animal and the river may have both been associated with speed. The connection between the two words is doubted in modern times, and they are likely to be Latin homonyms.[6]

Taxonomy

In 1758, Carl Linnaeus described the tiger in his work Systema Naturae and gave it the scientific name Felis tigris.[2] In 1929, the British taxonomist Reginald Innes Pocock subordinated the species under the genus Panthera using the scientific name Panthera tigris.[8][9]

Subspecies

Following Linnaeus's first descriptions of the species, several tiger zoological specimens were described and proposed as subspecies.[10] The validity of several tiger subspecies was questioned in 1999. Most putative subspecies described in the 19th and 20th centuries were distinguished on the basis of fur length and colouration, striping patterns and body size, hence characteristics that vary widely within populations. Morphologically, tigers from different regions vary little, and gene flow between populations in those regions is considered to have been possible during the Pleistocene. Therefore, it was proposed to recognize only two tiger subspecies as valid, namely P. t. tigris in mainland Asia, and P. t. sondaica in the Greater Sunda Islands. Mainland tigers are described as being larger in size with generally lighter fur and fewer stripes, while island tigers are smaller due to insular dwarfism, with darker coats and more numerous stripes.[11] The stripes of island tigers may break up into spotted patterns.[12]

This two-subspecies proposal was reaffirmed in 2015 by a comprehensive analysis of morphological, ecological and molecular traits of all putative tiger subspecies using a combined approach. The authors proposed recognition of only two subspecies, namely P. t. tigris comprising the Bengal, Malayan, Indochinese, South Chinese, Siberian and Caspian tiger populations of continental Asia, and P. t. sondaica comprising the Javan, Bali and Sumatran tiger populations of the Sunda Islands. The continental nominate subspecies P. t. tigris constitutes two clades: a northern clade composed of the Siberian and Caspian tiger populations, and a southern clade composed of all other mainland populations. The authors noted that this two-subspecies reclassification will impact tiger conservation management.[13] It would make captive breeding programs and future re-wilding of zoo-born tigers easier, as one tiger population could then be used to reinforce another. However, there is the risk that the loss of subspecies uniqueness could lead to less protection efforts for specific populations.[14]

In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group revised felid taxonomy in accordance with the two-subspecies proposal of the comprehensive 2015 study, and recognized the tiger populations in continental Asia as P. t. tigris, and those in the Sunda Islands as P. t. sondaica.[15] This two-subspecies view is still disputed by researchers, since the currently recognized six living subspecies can be distinguished genetically.[14] Results of a 2018 whole-genome sequencing of 32 samples support six monophyletic tiger clades corresponding with the six living proposed subspecies and indicate they descended from a common ancestor around 110,000 years ago.[16] Studies in 2021 and 2023 also affirmed the genetic distinctiveness and separation of these tigers.[17][18]

The following tables are based on the classification of the species Panthera tigris provided in Mammal Species of the World,[10] and also reflect the classification used by the Cat Classification Task Force in 2017:[15]

Panthera tigris tigris (Linnaeus, 1758)[2]
Populations Description Image
Bengal tiger This tiger inhabits the Indian subcontinent.[19] Linnaeus's scientific description of the tiger was based on descriptions by earlier naturalists such as Conrad Gessner and Ulisse Aldrovandi.[2] Bengal tiger skins in the collection of the Natural History Museum, London were described as bright orange-red with shorter fur and more spaced out stripes than northern-living tigers like the Siberian tiger.[9]
Caspian tiger formerly P. t. virgata (Illiger, 1815)[20] This population lived in west-central Asia, reach as far west as Turkey.[19] Illiger's description was not based on a particular specimen, but he only assumed that tigers in the Caspian area differ from those elsewhere.[20] It was later described having a bright rusty-red coat with thin and closely spaced brownish stripes,[21] and a broad occipital bone.[11] According to genetic analysis, it was closely related to the Siberian tiger.[22] It went extinct in the 1970s.[23]
Siberian tiger formerly P. t. altaica (Temminck, 1844)[24] The cat is found in the Russian Far East, Northeast China and possibly North Korea.[19] Temminck's description was based on an unspecified number of tiger skins with long hairs and dense coats that were traded between Korea and Japan. He assumed they originated in the Altai Mountains.[24] The Siberian tiger was later described as having pale coats with few dark brown stripes.[25] The lighter colouration may be due to longer exposure to sunlight during the summer. A Siberian tiger's coat becomes darker after molting.[11] The skull is described as shorter and broader then southern-living tigers.[26]
South China tiger formerly P. t. amoyensis (Hilzheimer, 1905)[27] This tiger historically lived in south-central China.[19] Hilzheimer's description was based on five tiger skulls purchased in Hankou, China. These skulls had shorter carnassials and molars than tigers from India with, a smaller cranium, orbits set closer together and larger postorbital processes. Skins of this tiger were described as being yellowish in colour with rhombus-like stripes.[27] It was noted to have a unique mtDNA haplotype.[15] This tiger may be extinct in wild as there has not been a confirmed sighting since the 1970s.[1]
Indochinese tiger formerly P. t. corbetti Mazák, 1968[28] The tiger is found on the Indochinese Peninsula.[19] Mazák's description was based on 25 specimens in museum collections that were smaller than tigers from India and had smaller skulls.[28] It was also said to have a darker coat than the Bengal tiger with more stripes; the stripes being narrower and having less "double stripes".[29]
Malayan tiger formerly P. t. jacksoni Luo et al., 2004[30] It was proposed as a distinct subspecies on the basis of mtDNA and micro-satellite sequences that differ from the Indochinese tiger.[30] In pelage colour or skull size, it does not differ significantly from Indochinese tigers.[29] There is no clear geographical barrier between tiger populations in northern Malaysia and southern Thailand.[1]
Panthera tigris sondaica (Temminck, 1844)[15]
Populations Description Image
Javan tiger formerly P. t. sondaica (Temminck, 1944)[24] Temminck based his description on an unspecified number of tiger skins with short and smooth hair.[24] Tigers from Java were small compared to tigers of the Asian mainland. The skull was relatively elongated and, compared to the Sumatran tiger, the stripes were longer, thinner and slightly greater in number.[29] The Javan tiger went extinct by the 1980s.[23]
Bali tiger formerly P. t. balica (Schwarz, 1912)[31] Schwarz based his description on a skin and a skull of an adult female tiger from Bali. He argued that its fur colour is brighter and its skull smaller than of tigers from Java.[31][32] A typical feature of Bali tiger skulls is the narrow occipital plane, which is similar to Javan tigers.[33] The tiger went extinct in the 1940s.[23]
Sumatran tiger formerly P. t. sumatrae Pocock, 1929[34] Pocock described a dark skin of a tiger from Sumatra as the type specimen.[34] It has broader and smaller nasal region than other island tigers[26][29] with many thick stripes.[29] This tiger has particularly long hairs around the face.[19]

Evolution

Two cladograms proposed for Panthera. The upper cladogram is based on two studies published in 2006 and 2009,[35][36] the lower one is based on studies published in 2010 and 2011.[37][38]

The tiger shares the genus Panthera with the lion, leopard, jaguar and snow leopard. Results of genetic analysis indicate that the tiger and snow leopard are sister species and about 2.88 million years ago, the tiger and the snow leopard lineages diverged from the other Panthera species.[35][39]

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 place it as basal to modern Panthera.[40][38] Panthera zdanskyi, which lived around the same time and place, was suggested to be a sister taxon of the modern tiger when it was examined in 2014.[38] However, as of 2023, at least two recent 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.[38] Middle to late Pleistocene tiger fossils were 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 were also found in Borneo and Palawan, Philippines.[43]

Results of a phylogeographic study indicate that all living tigers had a common ancestor 108,000 to 72,000 years ago.[30] A 2022 paleogenomic study of a Pleistocene tiger basal to living tigers concluded that modern tiger populations spread across Asia no earlier than 94,000 years ago. There is evidence of interbreeding between the lineage of modern mainland tigers and these ancient tigers.[44] The potential tiger range during the late Pleistocene and Holocene was predicted applying ecological niche modelling based on more than 500 tiger locality records combined with bioclimatic data. The resulting model shows a contiguous tiger range at the Last Glacial Maximum, indicating gene flow between tiger populations in mainland Asia. The tiger populations on the Sunda Islands and mainland Asia were possibly separated during interglacial periods.[45]

The tiger's full genome sequence was published in 2013. It was found to have repeat compositions much as other cat genomes and "an appreciably conserved synteny".[46]

Hybrids

Captive tigers were bred with lions to create hybrids called liger and tigon. The former born to a female tiger and male lion and the latter the result of a male tiger and female lion. They share physical and behavioural qualities of both parent species.[47] Because the lion sire passes on a growth-promoting gene, but the corresponding growth-inhibiting gene from the female tiger is absent, ligers grow far larger than either parent species. By contrast, the male tiger does not pass on a growth-promoting gene and the lioness passes on a growth inhibiting gene, hence tigons are around the same size as either species.[48] Breeding hybrids is now discouraged due to the emphasis on conservation.[47]

Characteristics

Siberian tiger in Aalborg Zoo, Denmark
Bengal tiger skeleton on display at the Museum of Osteology

The tiger is considered to be the largest living felid species.[12] However, there is some debate over averages compared to the lion. Since tiger populations vary greatly in size, the "average" size for a tiger may be less than a lion, while the biggest tigers are bigger than their lion counterparts.[43] The Siberian and Bengal tigers, along with the extinct Caspian are considered to be the largest of the species.[12] Bengal tigers average a total length of 3 m (9.8 ft), with males weighing 200–260 kg (440–570 lb) and females weighing 100–160 kg (220–350 lb).[49] Island tigers are the smallest, the Sumatran tigers have a total length of 2.2–2.5 m (7 ft 3 in – 8 ft 2 in) with a weight of 100–140 kg (220–310 lb) for males and 75–110 kg (165–243 lb) for females.[49] The extinct Bali tiger was even smaller.[12] It has been hypothesised that body sizes of different tiger populations may be correlated with climate and be explained by thermoregulation and Bergmann's rule.[12][11]

The tiger has a typical felid morphology. It has a muscular body with strong forelimbs, a large head and a tail that is about half the length of the rest of its body. There are five digits on the front feet and four on the back, all of which have retractable claws which are compact and curved. The ears are rounded, while the eyes have a round pupil.[12] The tiger's skull is large and robust, with a constricted front region, proportionally small, elliptical orbits, long nasal bones, and a lengthened cranium with a large sagittal crest.[50][12] It resembles a lion's skull; with the structure of the lower jaw and length of the nasals being the most reliable indicators for species identification.[50] The tiger has 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).[12][51] It has an average bite force at the canine tips of 1234.3 Newton.[52]

Coat

Tiger coat

Tiger fur tends to be short, except in the northern-living Siberian tiger. It has a mane-like heavy growth of fur around the neck and jaws and long whiskers, especially in males.[12] Its colouration is generally orange, but can vary from light yellow to dark red.[12][43][53] White fur covers the ventral surface, along with parts of the face.[12][54] It also has a prominent white spot on the back of their ears which are surrounded by black.[12] The tiger is marked with distinctive black or dark brown stripes; the patterns of which are unique in each individual.[12][55] The stripes are mostly vertical, but those on the limbs and forehead are horizonal. They are more concentrated towards the posterior and those on the trunk may or may not reach under the belly. The tips of stripes are generally sharp and some have gaps within them. Tail stripes are thick bands and a black tip marks the end.[56]

Stripes are likely advantageous for camouflage in vegetation with vertical patterns of light and shade, such as trees and long grass.[55] This is supported by a 1987 Fourier analysis study which concluded that the spatial frequencies of tiger stripes line up with their environment.[57] The tiger is one of only a few striped cat species; it is not known why spotted patterns and rosettes are the more common camouflage pattern among felids.[58] The orange colour may also aid in concealment as the tiger's prey are dichromats, and thus may perceive the cat as green and blended in with the vegetation.[59] The white dots on the ear may play a role in communication.[12]

Colour variations

Pseudo-melanistic white tiger

Three colour variantswhite, golden and nearly stripeless snow white 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 reddish-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 the white and wideband loci.[60] 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.[61]

Pseudo-melanistic tigers with thick, merged stripes have been recorded in Simlipal National Park and three Indian zoos; 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.[62]

Distribution and habitat

The tiger historically ranged from eastern Turkey and northern Afghanistan to Indochina, and from southeastern Siberia to Sumatra, Java and Bali.[12] As of 2022, it inhabits less than 7% of its historical distribution, and has a scattered range that includes the Indian subcontinent, the Indochinese Peninsula, Sumatra, the Russian Far East and northeastern China.[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).[63]

Global potential tiger habitat as of 2020[63]
Country Estimate
China 513,572 km2 (198,291 sq mi)
India 401,771 km2 (155,125 sq mi)
Myanmar 257,121 km2 (99,275 sq mi)
Russia 234,972 km2 (90,723 sq mi)
Indonesia 88,899 km2 (34,324 sq mi)
Laos 73,884 km2 (28,527 sq mi)
Cambodia 49,760 km2 (19,210 sq mi)
Thailand 46,884 km2 (18,102 sq mi)
Malaysia 41,713 km2 (16,105 sq mi)
Vietnam 37,187 km2 (14,358 sq mi)
Bhutan 26,905 km2 (10,388 sq mi)
Nepal 19,695 km2 (7,604 sq mi)
Bangladesh 8,897 km2 (3,435 sq mi)
North Korea 4,826 km2 (1,863 sq mi)

The tiger mainly lives in forest habitats and is highly adaptable.[49] Records in Central Asia indicate that it occurred foremost in Tugay riverine forests and inhabited hilly and lowland forests in the Caucasus.[64] In the Amur-Ussuri region, it inhabits Korean pine and temperate broadleaf and mixed forests; riparian forests serve as dispersal corridors, providing food and water for both tiger and ungulates.[65] On the Indian subcontinent, it inhabits mainly tropical and subtropical moist broadleaf forests, moist evergreen forests, tropical dry forests, alluvial plains and the mangrove forests of the Sundarbans.[66] In the Eastern Himalayas, tigers were documented in temperate forest up to an elevation of 4,200 m (13,800 ft) in Bhutan and of 3,630 m (11,910 ft) in the Mishmi Hills.[67][68] In Thailand, it lives in deciduous and evergreen forests.[69] In Sumatra, tigers range from lowland peat swamp forests to rugged montane forests.[70]

Behaviour and ecology

Tiger bathing in water

Camera trap data show that tigers in Chitwan National Park avoided locations frequented by people and were more active at night than by day.[71] In Sundarbans National Park, six radio-collared tigers were most active in the early morning with a peak around dawn and moved an average distance of 4.6 km (2.9 mi) per day.[72] A three-year long camera trap survey in Shuklaphanta National Park revealed that tigers were most active from dusk until midnight.[73] In northeastern China, tigers were crepuscular and active at night with activity peaking at dawn and at dusk; they exhibited a high temporal overlap with ungulate species.[74]

As with other felid species, tigers groom themselves, maintaining their coats by licking them and spreading oil from their sebaceous glands.[75] It will take to water, particularly on hot days. It is a powerful swimmer and easily transverses across rivers as wide as 8 km (5.0 mi).[55] Adults only occasionally climbs trees, but have been recorded climbing 10 m (33 ft) up a smooth pipal tree.[12] In general, tigers are less capable tree climbers than many other cats due to their size, but cubs under 16 months old may routinely do so.[76]

Social spacing

Adult tigers lead largely solitary lives. They establish and maintain home ranges, 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.[12][55] Two females in the Sundarbans had home ranges of 10.6 and 14.1 km2 (4.1 and 5.4 sq mi).[77] 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 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.[78] In Huai Kha Khaeng Wildlife Sanctuary, seven resident females had home ranges of 44.1–122.3 km2 (17.0–47.2 sq mi) and four resident males of 174.8–417.5 km2 (67.5–161.2 sq mi).[79] Four male problem tigers in Sumatra were translocated to national parks and needed 6–17 weeks to establish new home ranges of 37.5–188.1 km2 (14.5–72.6 sq mi).[80] Ten solitary females in Sikhote-Alin Biosphere Reserve had home ranges of 413.5 ± 77.6 km2 (159.7 ± 30.0 sq mi); when they had cubs of up to 4 months of age, their home ranges declined to 177.3 ± 53.5 km2 (68.5 ± 20.7 sq mi) and steadily grew to 403.3 ± 105.1 km2 (155.7 ± 40.6 sq mi) until the cubs were 13–18 months old.[81]

Tiger rubbing against tree to mark territory
Tiger rubbing against 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.[82] Young tigresses establish their first territories close to their mother's. Males, however, migrate further than their female counterparts and set out at a younger age to mark out their own area.[83] 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).[84] A young male may have to live as a transient in another male's territory until he is older and strong enough to challenge the resident male. Young males thus have an annual mortality rate of up to 35%. By contrast, young female tigers die at a rate of only around 5%.[83] Tigers mark their territories by spraying urine on vegetation and rocks, clawing or scent rubbing trees, and marking trails with feces, anal gland secretions and ground scrapings.[55][85][86][87] Scent markings also allow an individual to pick up information on another's identity. A tigress in oestrus will signal her availability by scent marking more frequently and increasing her vocalisations. Unclaimed territories, particularly those that belonged to a decreased individual, can be taken over in days or weeks.[55]

Male tigers are generally less tolerant of other males within their territories than females are of other females. Territory disputes are usually solved by intimidation rather than outright violence. Once dominance has been established, a male may tolerate a subordinate within his range, as long as they do not live in too close quarters. The most serious disputes tend to occur between two males competing for a female in oestrus.[88] Though tigers mostly live alone, relationships between individuals can be complex. Tigers are particularly social at kills, and a male tiger will share a carcass with the females and cubs within this territory and unlike male lions, will allow them to feed on the kill before he is finished with it. Though the female and male act amicably, females are more tense towards each other at a kill.[89][90]

Communication

Tiger baring teeth as a sign of aggression
Captive tiger roaring

During friendly encounters and bonding, tigers rub against each others' bodies.[91] Facial expressions include the "defense threat", which involves a wrinkled face, bared teeth, pulled-back ears, and widened pupils.[91][12] Both males and females show a flehmen response, a characteristic grimace, when sniffing urine markings. Males also use the flehman to detect the markings made by tigresses in oestrus.[12] Tigers 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.[92]

Tigers are normally silent but can produce numerous vocalisations.[93][94] 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. A tiger may roar three or four times in a row, and others may respond in kind. Tigers also roar during mating, and a mother will roar to call her cubs to her. When tense, tigers will 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.[12][95]

Aggressive encounters involve growling, snarling and hissing.[96] An explosive "coughing roar" or "coughing snarl" is emitted through an open mouth and exposed teeth.[12][96][97] Chuffing—soft, low-frequency snorting similar to purring in smaller cats—is heard in more friendly situations.[98] Mother tigers communicate with their cubs by grunting, while cubs call back with miaows.[99] A "woof" sound is produced when the animal is startled. It has also been recording emitting a deer-like "pok" sound for unknown reasons, but most often at kills.[100][101]

Hunting and diet

Tiger attacking a sambar deer in Ranthambore Tiger Reserve

The tiger is a carnivore and an apex predator feeding mainly on ungulates, with a particular preference for sambar deer, Manchurian wapiti, barasingha and wild boar. Tigers kill large prey like gaur,[102] but opportunistically kill much smaller prey like monkeys, peafowl and other ground-based birds, porcupines and fish.[12][55] Tiger attacks on adult Asian elephants and Indian rhinoceros have also been reported.[103][104][105] More often, tigers take the more vulnerable small calves.[106] When in close proximity to humans, tigers sometimes prey on domestic livestock and dogs.[12] Tigers occasionally consume vegetation, fruit and minerals for dietary fibre.[107]

Tigers learn to hunt from their mothers, which is important but not necessary for their success.[108] They usually hunt alone, but families hunt together when cubs are old enough.[109] A tiger travels up to 19.3 km (12.0 mi) per day in search of prey, using vision and hearing to find a target.[110] It also waits at a watering hole for prey to come by, particularly during hot summer days.[111][112] It is an ambush predator and when approaching potential prey, the tiger crouches, with head lowered, and hides in foliage. It switches between creeping forward and staying still. Tigers have been recorded dozing off while in still mode, and can stay in the same spot for as long as a day waiting for prey and launches an attack, when the prey is close enough,[113] usually within 30 m (98 ft).[49] If the prey spots it before then, the cat does not pursue further.[111] Tigers can sprint 56 km/h (35 mph) and leap 10 m (33 ft);[114][115] they are not long distance runners and give up a chase if prey outpaces them over a certain distance.[111]

Two tigers working together to kill an Indian 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. The tiger generally applies a bite to the throat until its target dies of strangulation.[12][116][117] Holding onto the throat puts the cat out of reach of the horns, antlers, tusks and hooves.[116][118] 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 to skull of a water buffalo.[119] They kill small prey with a bite to the back of the neck or skull.[120][49] Estimates of the success rate for hunting tigers ranges from a low 5% to a high of 50%. They are sometimes killed or injured by large or dangerous prey like gaur, buffalo and gaur.[49]

The tiger typically drags its kill for 183–549 m (600–1,801 ft) to a hidden, usually vegetated spot before eating. The tiger has the strength to drag the carcass of a fully grown buffalo for some distance, a feat three men struggle with. It rests for a while before eating and can consume as much as 50 kg (110 lb) of meat in one session, but feeds on a carcass for several days, leaving very little for scavengers.[121]

Enemies and competitors

An 1807 illustration of dholes attacking a tiger

Tigers may kill and even prey on other predators they coexist with.[122] In much of their range, tigers share habitat with leopards and dholes. They typically dominate both of them, though large packs of dholes can drive away a tiger,[123] or even kill it.[124] Tigers appear to inhabit the deep parts of a forest while these smaller predators are pushed closer to the fringes.[125] The three predators coexist by hunting different prey.[126] In one study, tigers were found to have killed prey that weighed an average of 91.5 kg (202 lb), in contrast to 37.6 kg (83 lb) for the leopard and 43.4 kg (96 lb) for the dhole.[127] Leopards can live successfully in tiger habitat when there is abundant food and vegetation cover, and there is no evidence of competitive exclusion.[126] Nevertheless, leopards avoid areas were tigers roam and are less common where tigers are numerous.[122][128][129]

Tigers tend to be wary of sloth bears, with their sharp claws, quickness and ability to stand on two legs. Tiger do sometimes prey on sloth bears by ambushing them when they are feeding at termite mounds.[130] Siberian tigers may attack, kill and prey on Ussuri brown and Ussuri black bears.[12] In turn, some studies show that brown bears frequently track down tigers to usurp their kills, with occasional fatal outcomes for the tiger.[131][132][133]

Reproduction and life cycle

Tiger family in Kanha Tiger Reserve

The tiger mates all year round, but most cubs are born between March and June, with another peak in September.[134] A tigress is in oestrus for three to six days, inbetween three to nine week intervals.[12] A resident male mates with all the females within his territory, who signal their receptiveness by roaring and marking.[135][136] Younger, transient males are also attracted, leading to a fight in which the more dominant male drives the usurper off.[134][135] 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 their tail to the side. Copulation is generally 20 to 25 seconds long, 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.[135] Tiger pairs may stay together for up to four days and mate multiple times.[137] Gestation ranges from 93 to 114 days, with an average of 103 to 105 days.[134]

A tigress gives birth in a secluded location, be it in dense vegetation, in a cave or under a rocky shelter.[138] Litters consist of as many as seven cubs, but two or three are more typical.[134][138] Newborn cubs weigh 785–1,610 g (27.7–56.8 oz), and are blind and altricial.[138] The mother licks and cleans her cubs, suckles them and viscously defends them from any potential threat.[134] She will only leave them alone to hunt, and even then does not travel far.[139] When a mother 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. The mortality rate for tiger cubs can reach 50% during these early months, causes of death include predators like dholes, leopards and pythons.[140] Young are able to see in a week, can leave the denning site in two months and around the same time they start eating meat.[134][141]

Mother tiger with 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, and she will guide them to the kill. Cubs bond though 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.[142] 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.[140] A cub can make a kill as early as 11 months, and reach independence around 18 to 24 months of age, males becoming independent earlier than females.[143] Radio-collared tigers in Chitwan started dispersing from their natal areas earliest at the age of 19 months.[84] Young females are sexual mature at three to four years, whereas males are at four to five years. Tigers may live up to 26 years.[12]

Tiger fathers play no role in raising the young, but he may encounter and interact with them. The resident male appears to visit the female-cub families within his territory. They may socalise and even share kills with them.[144][145] One male was recorded looking after cubs whose mother had died.[146] By defending his territory, the male is also protecting the females and cubs from other males.[147] When a new male takes over a territory, dependent cubs are at risk of being killed, as the male would want to sire his own young with the females. Older female cubs are tolerated but males may be treated as potential competitors.[148]

Threats

A hunting party poses with a killed Javan tiger, 1941

Major threats to the tiger include habitat destruction, habitat fragmentation and poaching for fur and body parts, which have simultaneously greatly reduced tiger populations in the wild.[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).[63] In the Tanintharyi Region of southern Myanmar, deforestation coupled with mining activities and a high hunting pressure threatens the tiger population in the area.[149] 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).[150] In Thailand, nine of 15 protected areas hosting tigers are isolated and fragmented offering a low probability for dispersal between them; and four of these do not harbour tigers any more at least since 2013.[151] In Peninsular Malaysia, an area of 8,315.7 km2 (3,210.7 sq mi) tiger habitat was cleared during 1988–2012, most of it for industrial plantations.[152] 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.[153] 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 southern China since 2001.[154][155]

Nam Et-Phou Louey National Park was considered the last important site for the tiger in Laos, but it has not been recorded since there at least since 2013; this population likely fell victim to indiscriminate snaring.[156] Tiger populations in India have been targetted by poachers since the 1990s and were extirpated in two tiger reserves in 2005 and 2009.[157]

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 and 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 were reported in India encompassing body parts of 893 tigers, and 403 seizures in Thailand involved mostly captive-bred tigers.[158]

Demand for tiger parts for use in traditional Chinese medicine has also been cited as a major threat to tiger populations.[159] 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.[160] 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.[161] Tiger bone glue is the prevailing tiger product purchased for medicinal purposes in Hanoi and Ho Chi Minh City.[162]

Conservation

Internationally, the tiger is protected under CITES Appendix I, banning trade of live tigers and their body parts.[1] In India, it has been protected since 1972 under Schedule I of the Wild Life (Protection) Act, 1972.[163] In 1973, the National Tiger Conservation Authority and Project Tiger were founded to gain public support for tiger conservation.[157] Since then, 53 tiger reserves covering an area of 75,796 km2 (29,265 sq mi) have been established in the country until 2022.[164] In Nepal, it has been protected since 1973 under the National Parks and Wildlife Conservation Act, 1973.[163] In Bhutan, it has been protected since 1969; the 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.[165] In Bangladesh, it has been protected since 1973 under the Wildlife (Preservation) Act and the Wildlife (Conservation and Security) Act, 2012.[161] In 2009, the Bangladesh Tiger Action Plan was initiated to stabilize the country's tiger population, maintain habitat and a sufficient prey base, improve law enforcement and cooperation between governmental agencies responsible for tiger conservation.[166] 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 of tiger killing and illegal trade of its body parts, and promoting public awareness through wildlife education programs.[167] In China, the trade in tiger body parts was banned in 1993, which helped to reduce the use of tiger bones in traditional Chinese medicine.[168]

Global wild tiger population
Country Year Estimate
India India 2023 3682–3925[169]
Russia Russia 2020 480–540[170]
Indonesia Indonesia 2016 400–600[171]
Bangladesh Bangladesh 2014 300–500[1]
Nepal Nepal 2022 355[172]
Thailand Thailand 2023 189[173]
Bhutan Bhutan 2023 131[174]
Malaysia Malaysia 2022 <150[175]
China China 2018 55[176]
Myanmar Myanmar 2018 22[177]
Total 5,764–6,467

In the 1990s, a new approach to tiger conservation was developed: Tiger Conservation Units (TCUs), which are blocks of habitat that have the potential to host tiger populations in 15 habitat types within five bioregions. Altogether 143 TCUs were identified and prioritized based on size and integrity of habitat, poaching pressure and population status. They range in size from 33 to 155,829 km2 (13 to 60,166 sq mi).[66]

In the 1940s, the tiger was on the brink of extinction in Russia. Anti-poaching controls were initialised, and a network of protected areas, so-called zapovedniks, were instituted, leading to a rise in the tiger population.[178] Increases in patrol effort during 2011–2014 in four protected areas contributed to reducing poaching, stabilising the tiger population and improving protection of ungulate populations.[179]

In 1994, the Indonesian Sumatran Tiger Conservation Strategy addressed the potential crisis that tigers faced in Sumatra. The Sumatran Tiger Project (STP) was initiated in June 1995 in and around the Way Kambas National Park to ensure the long-term viability of wild Sumatran tigers and to accumulate data on tiger life-history characteristics vital for the management of wild populations.[180] By August 1999, the teams of the STP had evaluated 52 sites of potential tiger habitat in Lampung Province, of which only 15 these were intact enough to contain tigers.[181] In the framework of the STP a community-based conservation program was initiated to document the tiger-human dimension in the park to enable conservation authorities to resolve tiger-human conflicts based on a comprehensive database rather than anecdotes and opinions.[182]

The Wildlife Conservation Society and Panthera Corporation formed the collaboration Tigers Forever, with field sites including the world's largest tiger reserve, the 21,756 km2 (8,400 sq mi) Hukaung Valley in Myanmar. Other reserves were located in India, Thailand, Laos, Cambodia and the Russian Far East covering in total about 260,000 km2 (100,000 sq mi).[183]

Tigers have been studied in the wild using a variety of techniques. Tiger population have been estimated using plaster casts of their pugmarks, although this method was criticized as being inaccurate.[184] More recent techniques include the use of camera traps and studies of DNA from tiger scat, while radio-collaring has been used to track tigers in the wild.[185] Tiger spray has been found to be just as good, or better, as a source of DNA than scat.[186]

Relationship with humans

Hunting

Tiger hunting on elephant-back in India, 1808

A tiger hunt is painted on the Bhimbetka rock shelters in India and dated to 5,000–6,000 years ago. Thousands of years later, Emperor Samudragupta was depicted slaying tigers on coins. Tiger hunting became an established sport under the Mughal Empire in the 16th century. The cats were chased on horseback and killed with spears. Emperor Akbar participated in such activities and one of his hunts is the subject of a painting from the Akbarnama. Following Akbar, Emperor Jahangir will introduce musket to tiger hunts and eventually, elephant would be ridden. The British East India Company would pay for bounties on tigers as early as 1757 and tiger hunting would continue under British Raj.[187] Tiger killings were particularly high in the 19th and early 20th centuries; as an estimated 80,000 cats were killed between 1875 and 1925.[188][189] King George V on his visit to Colonial India in 1911 killed 39 tigers in a matter of 10 days.[190]

Attacks

A tiger in the Sundarbans. This area is known for its relatively high level of tiger attacks.

Tigers are said to have directly killed more people than any other wild mammal.[191] In most areas, the big cats typically avoid humans, but attacks are a risk wherever people coexist with them.[192][193] Dangerous encounters are more likely to occur in edge habitats, between wild and agricultural areas.[192] Most attacks on humans are defensive, including protection of young. However, tigers do sometimes see people as prey.[193] They hunt people the same way they hunt other prey, by ambush and with a killing bite to the neck. A tiger inflicted wound also carries the risk of infection.[192] Man-eating tigers tend to be old and disabled.[55] Those they have been driven from their home ranges and territories are also at risk of turning to man-eating.[194]

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 famed hunter Jim Corbett.[195] Corbett recorded that the tigress suffered from broken teeth and thus unable to kill normal prey. Modern authors speculate that feeding on meagre human flesh forced the cat to kill more and more.[196] Tiger attacks were particularly high in Singapore during the mid-19th century, when plantations expanded into the animal's habitat.[197] The number of deaths ranged from 200 to 300 annually in the 1840s.[198]

Tiger predation on humans is highest in the Sundarbans. An estimated 129 people were killed between 1969 and 1971. In the 10 years prior to that period, about 100 attacks per year in the Sundarbans.[191] Victims of tigers 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. These tools have been credited with reducing tiger attacks to only 22 per year in the 1980s.[199] Because of rapid habitat loss attributed to climate change, tiger attacks have increased in the Sundarbans in the 21st century.[200]

In captivity

Tiger at Big Cat Rescue in 2014
Publicity 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 amphitheaters; they were slaughtered in hunts and used for public executions of criminals. Mongol Emperor Kublai Khan is reported to have kept tigers in the 13th century. Starting in the Middle Ages, tigers were being kept in European menageries. In 1830, two tigers and a lion were accidentally put in the same exhibit at the Tower of London. This lead to a fight between them and, after they were separated, the lion died of its wounds.[201] 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.[202]

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 big cats 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. Tiger trainer Sara Houckle was dubbed "the Tiger Whisperer", as she trained the cats to obey her by whispering to them.[203] Siegfried & Roy became famous for performing with white tigers in Las Vegas. The act ended in 2003 when a tiger named Mantacore attacked Roy during a performance.[204] The use of tigers and other animals in shows would eventually decline 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 would restrict or ban such acts.[205] According to a 2009 analysis, tigers were the most traded circus animals.[206]

Tigers have become popular in the exotic pet trade, particularly in the United States.[207] The World Wide Fund for Nature (WWF) estimated that in the US, 5,000 tigers were kept in captivity in 2020, with only 6% of them being in zoos and other facilities approved by the Association of Zoos and Aquariums. The WWF argues that private collectors are 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.[208] The keeping of tigers and other big cats by private individuals was banned in the US in 2022 under the Big Cat Public Safety Act. Those who owned big cats at the time of the signing were expected to register with the United States Fish and Wildlife Service before 18 June 2023.[209] The WWF also estimated in 2020 that 7,000–8,000 tigers were held in "tiger farm" facilities in China and Southeast Asia. These tigers are bred to be used for traditional medicine and appear to pose a threat to wild populations by rising demand for tiger parts.[208]

Cultural significance

Tiger-shaped bronze from Zhou-era China, (c. 900 BC)

The tiger is among the most famous of charismatic megafauna. It has been labelled as "a rare combination of courage, ferocity and brilliant colour".[134] In a 2004 online poll conducted by cable television channel Animal Planet, involving more than 50,000 viewers from 73 countries, the tiger was voted the world's favourite animal with 21% of the vote, narrowly beating the dog.[210] Likewise, a 2018 study found the tiger to be the most popular wild animal based on surveys, as well as appearances on websites of major zoos and posters of some animated movies.[211]

While the lion represented royalty and power in Western culture, the tiger filled such a role in Asia. In ancient China, the tiger was seen as the "king of the forest" and symbolised the power of the emperor.[212] In Chinese astrology, the tiger is the third out of 12 symbols in the zodiac and controls the period of the day between 3 am and 5 am. The Year of the Tiger is thought to bring "dramatic and extreme events". The White Tiger is one of the Four Symbols of the Chinese constellations, representing the west along with the yin and the season of autumn. It is the counterpart to the Azure Dragon, which conversely symbolises the east, yang and springtime.[213] The tiger is one of the animals displayed on the Pashupati seal of the Indus Valley civilisation. The big cat was depicted on seals and coins during the Chola Dynasty of southern India, as it was the official emblem.[214]

The Hindu goddess Durga riding a tiger. Guler school, early 18th century

Tigers have had religious significance, even being worshiped. In Buddhism, the tiger, monkey and deer are Three Senseless Creatures, the tiger symbolising anger.[215] In Bhutan, the tiger is venerated as one of the four powerful animals called the "four dignities", and a tigress is believed to have carried Padmasambhava from Singye Dzong to the Paro Taktsang monastery in the late 8th century.[165] In Hinduism, the tiger is the vehicle for the goddess of feminine power and peace, Durga, whom the gods created to fight demons. Similarly, in the Greco-Roman world, the tiger was depicted being ridden by the god Dionysus. In Korean mythology, tigers are messengers of the Mountain Gods.[216] The Warli of western India worship the tiger-like god Waghoba. The Warli believe that shrines and sacrifices to the deity will lead to better coexistence with the local big cats, both tigers and leopards, and that Waghoba will protect them when they enter the forests.[217]

In both Chinese and Korean culture, tigers are seen as a protectors against evil spirits, and their image was used to decorate homes and tombs.[212][218] In the folklore of Malaysia and Indonesia, "tiger shamans" heal the sick by evoking the big cat. People turning into tigers and the inverse has also been widespread, in particular weretigers are people who could change into tigers and back again. The Mnong people of Indochina believed that tigers could transform into humans.[219] Among some indigenous peoples of Siberia, it was believed that men would seduce women by transforming into tigers.[212]

Blake's original printing of The Tyger, 1794

The tiger's cultural reputation is generally that of a fierce and powerful animal. William Blake's 1794 poem "The Tyger" portrays the animal as the duality of beauty and ferocity. It is the sister poem to "The Lamb" in Blake's Songs of Innocence and of Experience and he ponders why God would create such different creatures. The tiger is featured in the medieval Chinese novel Water Margin, where the cat battles and is slain by the bandit Wu Song, while the tiger Shere Khan in Rudyard Kipling's 1894 The Jungle Book is the mortal enemy of the human protagonist Mowgli. The image of the friendly tame tiger has also existed in culture, notably Tigger, the Winnie-the-Pooh character and Tony the Tiger, the Kellogg's cereal mascot.[220]

See also

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Bibliography

External links