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Lion

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Not to be confused with Mountain lion.

This article is about the animal. For Other uses, see Lion (disambiguation).

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 hairy tuft at the end 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 large ungulates. The lion is an apex and keystone predator; although some lions scavenge when opportunities occur and have been known to hunt humans, lions typically do not actively seek out and prey on humans.

Lion

Temporal range: Pleistocene–Present

PreꞒ

Ꞓ

↓

Male in Okonjima, NamibiaFemale (lioness) in Okonjima

Conservation status

Vulnerable (IUCN 3.1)[2]

CITES Appendix II (CITES)[a][2]

Scientific classificationDomain:EukaryotaKingdom:AnimaliaPhylum:ChordataClass:MammaliaOrder:CarnivoraSuborder:FeliformiaFamily:FelidaeSubfamily:PantherinaeGenus:PantheraSpecies:

P. leo[1]

Binomial namePanthera leo[1]

(Linnaeus, 1758)[3]

SubspeciesP. l. leoP. l. melanochaita†P. l. sinhaleyusHistorical and present distribution of the lion in Africa, Asia and Europe

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]

Taxonomy

The 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]

Subspecies

Range 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 records

Skull 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.[29] It arose when a cave lion population in Beringia became isolated south of the Cordilleran Ice Sheet about 370,000 years ago.[30][31] A fossil from Edmonton dates to 11,355 ± 55 years ago.[32]

Evolution

red 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][33][34] 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.[34] The lion-leopard clade was distributed in the Asian and African Palearctic since at least the early Pliocene.[35] The earliest fossils recognisable as lions were found at Olduvai Gorge in Tanzania and are estimated to be up to 2 million years old.[33]

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][36][37] 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.[38] 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.[38]

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.[39] 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]

Hybrids

Further information: Panthera hybrid

In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose.[40][41] 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.[42][43] The leopon is a hybrid between a lion and leopard.[44]

Description

A 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.[45][46] 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" that is formed from the final, fused sections of tail bone. 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]

Size

Among felids, the lion is second only to the tiger in size.[46] The size and weight of adult lions vary across its range and habitats.[52][53][54][55] Accounts of a few individuals that were larger than average exist from Africa and India.[45][56][57][58]

AverageFemale lionsMale lionsHead-and-body length160–184 cm (63–72 in)[59]184–208 cm (72–82 in)[59]Tail length72–89.5 cm (28.3–35.2 in)[59]82.5–93.5 cm (32.5–36.8 in)[59]Weight118.37–143.52 kg (261.0–316.4 lb) in Southern Africa,[52]

119.5 kg (263 lb) in East Africa,[52]

110–120 kg (240–260 lb) in India[53]186.55–225 kg (411.3–496.0 lb) in Southern Africa,[52]

174.9 kg (386 lb) in East Africa,[52]

160–190 kg (350–420 lb) in India[53]

Mane

A six-year-old male in Phinda Private Game Reserve

Male in Pendjari National Park

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.[60] 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.[46] It starts growing when lions enter adolescence, when testosterone levels increase, and reach their full size at around four years old.[61] Cool ambient temperatures in European and North American zoos may result in a heavier mane.[62] On average, Asiatic lions have sparser manes than African lions.[63]

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.[64][65] Unlike in other felid species, female lions consistently interact with multiple males at once.[66] Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed.[67][68] 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.[69]

Almost all male lions in Pendjari National Park are either maneless or have very short manes.[70] Maneless lions have also been reported in Senegal, in Sudan's Dinder National Park and in Tsavo East National Park, Kenya.[71] Castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production.[72]

Rarely, lionesses (in the wild and in captivity) have been observed to grow manes.[73][74] Increased testosterone may be the cause of maned lionesses reported in northern Botswana.[75]

Colour variation

Further 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.[76] White lions are selected for breeding in captivity.[77] They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts.[78]

Distribution and habitat

Lion in Gir National Park

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.[45] 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.[79] 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.[80] In the 1960s, it became extinct in North Africa, except in the southern part of Sudan.[81][79][82]

In southern Europe and Asia, the lion once ranged in regions where climatic conditions supported an abundance of prey.[83] In Ukraine, the modern lion was present from about 6,400 to 2,000 years BP.[84] In Greece, it was common, as reported by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100.[45] 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.[85] The last live lion in Iran was sighted in 1942, about 65 km (40 mi) northwest of Dezful,[86] although the corpse of a lioness was found on the banks of the Karun river in Khuzestan province in 1944.[87] It once ranged from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India.[88]

Behaviour and ecology

Lions spend much of their time resting; they are inactive for about twenty hours per day.[89] 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.[90]

Group organisation

Lion pride in Etosha National Park

A 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".[91] Females form the stable social unit in a pride and do not tolerate outside females.[92] The majority of females remain in their birth prides while all males and some females will disperse.[93] 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.[94] The sole exception to this pattern is the Tsavo lion pride that always has just one adult male.[95] Prides act as fission–fusion societies, and members will split into subgroups that keep in contact with roars.[96]

Nomadic lions range widely and move around sporadically, either in pairs or alone.[91] Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.[97] Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them.[98] Males spend years in a nomadic phase before gaining residence in a pride.[99] A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.[100] 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.[101]

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).[102][103] The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".[91] Males associated with a pride patrol the fringes.[46] 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 defense against intruders, while others lag behind.[104] Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.[105] 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.[106] The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.[97] Dominance hierarchies do not appear to exist among individuals of either sex in a pride.[107]

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.[108] 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.[109]

Hunting and diet

Male lion and cub feeding on a Cape buffalo in Sabi Sand Game Reserve

Four lionesses catching a buffalo in the Serengeti

A skeletal mount of a lion attacking a common eland, on display at The Museum of Osteology

The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.[110][111] Its prey consists mainly of ungulates, particularly blue wildebeest, plains zebra, African buffalo, common warthog, gemsbok and giraffe.[112] In India, chital and sambar deer are the most common wild prey,[46][112][113] while livestock contributes significantly to lion kills outside protected areas.[114] They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus and small prey like dik-dik, hyraxes, hares and monkeys.[112][115] Unusual prey include porcupines and small reptiles. Lions kill other predators but seldom consume them.[116]

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.[117] Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.[97] 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.[118][119] In typical 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.[120] Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.[121] 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.[122]

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.[123] 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.[124] One study in 2018 recorded a lion running at a top speed of 74.1 km/h (46.0 mph).[125] They take advantage of factors that reduce visibility; many kills take place near some form of cover or at night.[126] 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.[127] It has a bite force of 1314.7 Newton at the canine tip and 2023.7 Newton at the carnassial notch.[128]

Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover.[129] 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.[97] Large kills are shared more widely among pride members.[130] An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).[131] Lions gorge themselves and eat up to 30 kg (66 lb) in one session.[87] 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.[129] Lions defend their kills from scavengers such as vultures and hyenas.[97]

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.[132] Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions.[55] Carrion is thought to provide a large part of lion diet.[133]

Predatory competition

Lioness chasing a spotted hyena in Kruger National Park

Lioness stealing a kill from a leopard in Kruger National Park

Lions and spotted hyenas occupy a similar ecological niche and compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%.[134] Lions typically ignore hyenas unless they are on a kill or are being harassed, while the latter tend to visibly react to the presence of lions with or without the presence of food. In the Ngorongoro crater, lions subsist largely on kills stolen from hyenas, causing them to increase their kill rate.[135] In Botswana's Chobe National Park, the situation is reversed as hyenas there frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills.[136] When confronted on a kill, hyenas may either leave or wait patiently at a distance of 30–100 m (98–328 ft) until the lions have finished.[137] Hyenas may feed alongside lions and force them off a kill. The two species attack one another even when there is no food involved for no apparent reason.[138] Lions can account for up to 71% of hyena deaths in Etosha National Park. Hyenas have adapted by frequently mobbing lions that enter their home ranges.[139] When the lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly.[140]

Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance.[141] Cheetahs often lose their kills to lions or other predators.[142] A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.[143] Cheetahs avoid their competitors by hunting at different times and habitats.[144] Leopards, by contrast, do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around the same time of day. In addition, there is no evidence that lions effect leopard abundance.[145] Leopards take refuge in trees, though lionesses occasionally attempt to climb up and retrieve their kills.[146]

Lions similarly dominate African wild dogs, taking their kills and dispatching pups or adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.[147] However, there are a few reported cases of old and wounded lions falling prey to wild dogs.[148][149]

Reproduction and life cycle

Lions mating at Masai Mara

A lion cub in Masai Mara

Most lionesses reproduce by the time they are four years of age.[150] Lions do not mate at a specific time of year and the females are polyestrous.[151] Like those of other cats, the male lion's penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation.[152][153] A lioness may mate with more than one male when she is in heat.[154] Lions of both sexes may be involved in group homosexual and courtship activities. Males will also head-rub and roll around with each other before mounting each other.[155][156] Generation length of the lion is about seven years.[157] The average gestation period is around 110 days;[151] the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den.[158] Lion cubs are born blind, their eyes opening around seven days after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.[159] To avoid a buildup of scent attracting the attention of predators, the lioness moves her cubs to a new den site several times a month, carrying them one-by-one by the nape of the neck.[158]

Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.[158] Sometimes the introduction to pride life occurs earlier, particularly if other lionesses have given birth at about the same time.[97][160] When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults other than their mother. They soon begin to immerse themselves in the pride life, however, playing among themselves or attempting to initiate play with the adults.[160] Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs varies—one male could patiently let the cubs play with his tail or his mane, while another may snarl and bat the cubs away.[161]

Video of a lioness and her cubs in Phinda Reserve

Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride. The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival, and sucklings are not dominated by older cubs.[97][160] Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest.[162]

When one or more new males oust the previous males associated with a pride, the victors often kill any existing young cubs, perhaps because females do not become fertile and receptive until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male.[163] Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs. Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age,[164] while some females may leave when they reach the age of two.[93] When a new male lion takes over a pride, adolescents both male and female may be evicted.[165]

Health and mortality

Lions in a tree near Lake Nakuru

Lions may live 12–17 years in the wild.[46] Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.[166] Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill.[167] Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey.[168] Nile crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in crocodile stomachs.[169]

Ticks commonly infest the ears, neck and groin regions of the lions.[170][171] Adult forms of several tapeworm species of the genus Taenia have been isolated from lion intestines, having been ingested as larvae in antelope meat.[172] Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962, resulting in lions becoming emaciated and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals.[173] A more recent outbreak in 2001 killed six lions.[174]

Captive lions have been infected with canine distemper virus (CDV) since at least the mid-1970s.[175] CDV is spread by domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and encephalitis.[176] Feline immunodeficiency virus and lentivirus also affect captive lions.[177][178]

Communication

Head rubbing among pride members is a common social behaviour.

When resting, lion socialisation occurs through a number of behaviours; the animal's expressive movements are highly developed. The most common peaceful, tactile gestures are head rubbing and social licking,[179] which have been compared with the role of allogrooming among primates.[180] Head rubbing, nuzzling the forehead, face and neck against another lion appears to be a form of greeting[181] and is seen often after an animal has been apart from others or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.[182] Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure. The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves.[183]

Lion roar

Duration: 10 seconds.0:10

A captive lion roaring

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Lions have an array of facial expressions and body postures that serve as visual gestures.[184] A common facial expression is the "grimace face" or flehmen response, which a lion makes when sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled nose, closed eyes and relaxed ears.[185] Lions also use chemical and visual marking;[184] males spray urine[186] and scrape plots of ground and objects within the territory.[184]

The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of growling, snarling, meowing and roaring. Other sounds produced include purring, puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of 8 kilometres (5 mi).[187] They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into grunts.[188][189]

Conservation

The lion is listed as Vulnerable on the IUCN Red List. The Indian population is listed on CITES Appendix I and the African population on CITES Appendix II.[2]

In Africa

Video of a wild lioness

Several large and well-managed protected areas in Africa host large lion populations. Where an infrastructure for wildlife tourism has been developed, cash revenue for park management and local communities is a strong incentive for lion conservation.[2] Most lions now live in East and Southern Africa; their numbers are rapidly decreasing, and fell by an estimated 30–50% in the late half of the 20th century. Primary causes of the decline include disease and human interference.[2] In 1975, it was estimated that since the 1950s, lion numbers had decreased by half to 200,000 or fewer.[190] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004.[191][81]

In the Republic of the Congo, Odzala-Kokoua National Park was considered a lion stronghold in the 1990s. By 2014, no lions were recorded in the protected area so the population is considered locally extinct.[192] The West African lion population is isolated from the one in Central Africa, with little or no exchange of breeding individuals. In 2015, it was estimated that this population consists of about 400 animals, including fewer than 250 mature individuals. They persist in three protected areas in the region, mostly in one population in the W A P protected area complex, shared by Benin, Burkina Faso and Niger. This population is listed as Critically Endangered.[13] Field surveys in the WAP ecosystem revealed that lion occupancy is lowest in the W National Park, and higher in areas with permanent staff and thus better protection.[193]

A population occurs in Cameroon's Waza National Park, where between approximately 14 and 21 animals persisted as of 2009.[194] In addition, 50 to 150 lions are estimated to be present in Burkina Faso's Arly-Singou ecosystem.[195] In 2015, an adult male lion and a female lion were sighted in Ghana's Mole National Park. These were the first sightings of lions in the country in 39 years.[196] In the same year, a population of up to 200 lions that was previously thought to have been extirpated was filmed in the Alatash National Park, Ethiopia, close to the Sudanese border.[197][198]

In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild prey base for lions, reduce factors that lead to further fragmentation of populations, and make lion–human coexistence sustainable.[199][200] Lion depredation on livestock is significantly reduced in areas where herders keep livestock in improved enclosures. Such measures contribute to mitigating human–lion conflict.[201]

In Asia

A lioness in Gir National Park

The last refuge of the Asiatic lion population is the 1,412 km2 (545 sq mi) Gir National Park and surrounding areas in the region of Saurashtra or Kathiawar Peninsula in Gujarat State, India. The population has risen from approximately 180 lions in 1974 to about 400 in 2010.[202] It is geographically isolated, which can lead to inbreeding and reduced genetic diversity. Since 2008, the Asiatic lion has been listed as Endangered on the IUCN Red List.[12] By 2015, the population had grown to 523 individuals inhabiting an area of 7,000 km2 (2,700 sq mi) in Saurashtra.[203][204][205] In 2017, about 650 individuals were recorded during the Asiatic Lion Census.[206]

The presence of numerous human settlements close to Gir National Park resulted in conflict between lions, local people and their livestock.[207][203] Some consider the presence of lions a benefit, as they keep populations of crop damaging herbivores in check.[208]

Captive breeding

Two captive male Asiatic lions in Sanjay Gandhi National Park, India

Lions imported to Europe before the middle of the 19th century were possibly foremost Barbary lions from North Africa, or Cape lions from Southern Africa.[209] Another 11 animals thought to be Barbary lions kept in Addis Ababa Zoo are descendants of animals owned by Emperor Haile Selassie. WildLink International in collaboration with Oxford University launched an ambitious International Barbary Lion Project with the aim of identifying and breeding Barbary lions in captivity for eventual reintroduction into a national park in the Atlas Mountains of Morocco.[210] However, a genetic analysis showed that the captive lions at Addis Ababa Zoo were not Barbary lions, but rather closely related to wild lions in Chad and Cameroon.[211]

In 1982, the Association of Zoos and Aquariums started a Species Survival Plan for the Asiatic lion to increase its chances of survival. In 1987, it was found that most lions in North American zoos were hybrids between African and Asiatic lions.[212] Breeding programs need to note origins of the participating animals to avoid cross-breeding different subspecies and thus reducing their conservation value.[213] Captive breeding of lions was halted to eliminate individuals of unknown origin and pedigree. Wild-born lions were imported to American zoos from Africa between 1989 and 1995. Breeding was continued in 1998 in the frame of an African lion Species Survival Plan.[214]

About 77% of the captive lions registered in the International Species Information System in 2006 were of unknown origin; these animals might have carried genes that are extinct in the wild and may therefore be important to the maintenance of the overall genetic variability of the lion.[62]

Interactions with humans

In zoos and circuses

Lion at Melbourne Zoo

19th-century etching of a lion tamer in a cage with lions and tigers

Lions are part of a group of exotic animals that have been central to zoo exhibits since the late 18th century. Although many modern zoos are more selective about their exhibits,[215] there are more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They are considered an ambassador species and are kept for tourism, education and conservation purposes.[216] Lions can live over twenty years in captivity; for example, three sibling lions at the Honolulu Zoo lived to the age of 22 in 2007.[217][218]

The first European "zoos" spread among noble and royal families in the 13th century, and until the 17th century were called seraglios. At that time, they came to be called menageries, an extension of the cabinet of curiosities. They spread from France and Italy during the Renaissance to the rest of Europe.[219] In England, although the seraglio tradition was less developed, lions were kept at the Tower of London in a seraglio established by King John in the 13th century;[220][221] this was probably stocked with animals from an earlier menagerie started in 1125 by Henry I at his hunting lodge in Woodstock, Oxfordshire, where according to William of Malmesbury lions had been stocked.[222]

Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with roomier cages was built in the 1870s.[223] Further changes took place in the early 20th century when Carl Hagenbeck designed enclosures with concrete "rocks", more open space and a moat instead of bars, more closely resembling a natural habitat. Hagenbeck designed lion enclosures for both Melbourne Zoo and Sydney's Taronga Zoo; although his designs were popular, the use of bars and caged enclosures prevailed in many zoos until the 1960s.[224] In the late 20th century, larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered dens allowed visitors to come closer than ever to the animals; some attractions such as the Cat Forest/Lion Overlook of Oklahoma City Zoological Park placed the den on ground level, higher than visitors.[225]

Lion taming has been part of both established circuses and individual acts such as Siegfried & Roy. The practice began in the early 19th century by Frenchman Henri Martin and American Isaac Van Amburgh, who both toured widely and whose techniques were copied by a number of followers.[226] Van Amburgh performed before Queen Victoria in 1838 when he toured Great Britain. Martin composed a pantomime titled Les Lions de Mysore ("the lions of Mysore"), an idea Amburgh quickly borrowed. These acts eclipsed equestrianism acts as the central display of circus shows and entered public consciousness in the early 20th century with cinema. In demonstrating the superiority of human over animal, lion taming served a purpose similar to animal fights of previous centuries.[226] The ultimate proof of a tamer's dominance and control over a lion is demonstrated by the placing of the tamer's head in the lion's mouth. The now-iconic lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[227]

Hunting and games

Main article: Lion hunting

See also

List of largest catsMapogo lion coalitionRoar (film) from 1981

Notes

^ Populations of India are listed in Appendix I.

References

External links

Look up lion in Wiktionary, the free dictionary.

Wikispecies has information related to Lion.

Wikisource has the text of the 1911 Encyclopædia Britannica article "Lion".

Media related to Lion at Wikimedia CommonsIUCN/SSC Cat Specialist Group. "Lion Panthera leo"."Lion Conservation Fund".The Portugal News (2014). "Rare desert lion killed in Angola after supplying unprecedented data". Archived from the original on 2 August 2018. Retrieved 24 May 2018."Lion" . Collier's New Encyclopedia. 1921.

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Tiger

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"Tigress" redirects here. For other uses, see Tiger (disambiguation) and Tigress (disambiguation).

The tiger (Panthera tigris) is the largest living cat species 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 island tigers of the Sunda Islands.

Tiger

Temporal range: Early Pleistocene – Present

PreꞒ

Ꞓ

↓

A Bengal tigress in Kanha Tiger Reserve, India

Conservation status

Endangered (IUCN 3.1)[1]

CITES Appendix I (CITES)[1]

Scientific classificationDomain:EukaryotaKingdom:AnimaliaPhylum:ChordataClass:MammaliaOrder:CarnivoraSuborder:FeliformiaFamily:FelidaeSubfamily:PantherinaeGenus:PantheraSpecies:

P. tigris

Binomial namePanthera tigris

(Linnaeus, 1758)[2]

SubspeciesP. t. tigrisP. t. sondaica†P. t. acutidens†P. t. soloensis†P. t. trinilensisTiger distribution as of 2022Synonyms[3]Felis tigris Linnaeus, 1758Tigris striatus Severtzov, 1858Tigris regalis Gray, 1867

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 such as deer and wild boar, which it takes by ambush. It lives a mostly solitary life and occupies home ranges, which it defends 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 reasons for this decline are habitat destruction and fragmentation due to deforestation, poaching for fur, and the illegal trade of tiger body parts for medicinal purposes. Tigers are also victims of human–wildlife conflict for attacking and preying on livestock in areas where natural prey is scarce. The species is legally protected in all range countries, which have ratified conservation action plans, established anti-poaching patrols and schemes for monitoring tiger populations.

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 Classical Greek τίγρις 'tigris'.[4] In the 1st century BC, Roman scholar Marcus Terentius Varro argued that the word tigris originated from the Armenian language and means 'arrow', which is also the name of the fast-flowing river Tigris.[5] The Middle Persian tigr also has both meanings; however, today, the connection between 'arrow' and the river's name is doubted, 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; his scientific description was based on descriptions by earlier naturalists such as Conrad Gessner and Ulisse Aldrovandi.[2] In 1929, the British taxonomist Reginald Innes Pocock subordinated the species under the genus Panthera, using the scientific name Panthera tigris.[7][8]

Subspecies

Following Linnaeus's first descriptions of the species, nine recent tiger subspecies have been proposed between the early 19th and early 21st centuries, 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; since fur characteristics vary widely within populations, 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. 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.[10] In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group revised felid taxonomy 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 monophyletic 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 are yellowish with rhombus-like stripes.[26] It has a unique mtDNA haplotype.[12] 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)[27]This tiger population occurs on the Indochinese Peninsula.[17] Indochinese tiger specimens are smaller with smaller skulls than specimens from India.[27] They appear to have darker fur than specimens from India with stripes being slightly narrower.[28] Malayan tiger formerly P. t. jacksoni (Luo et al., 2004)[29]The Malayan tiger was proposed as a distinct subspecies on the basis of mtDNA and micro-satellite sequences that differ from the Indochinese tiger.[29] It does not differ significantly in fur colour or skull size from Indochinese tigers.[28] 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. The skull was relatively elongated and, compared to the Sumatran tiger, the stripes were longer, thinner and slightly greater in number.[28] The Javan tiger is thought to have gone extinct by the 1980s.[23] †Bali tiger formerly P. t. balica (Schwarz, 1912)[30]This tiger occurred on Bali and had a brighter fur and a smaller skull than the Javan tiger.[30][31] A typical feature of Bali tiger skulls is the narrow occipital bone, which is similar to the Javan tiger's skull.[32] The tiger went extinct in the 1940s.[23] Sumatran tiger formerly P. t. sumatrae (Pocock, 1929)[33]The type specimen from Sumatra had dark fur.[33] The Sumatran tiger has particularly long hair around the face,[17] thick body stripes,[28], and a broader and smaller nasal bone than other island tigers.[28][19]

Evolution

Two cladograms proposed for Panthera. The upper cladogram is based on two studies published in 2006 and 2009,[34][35] the lower one is based on studies published in 2010 and 2011.[36][37]

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 whose lineages split from each other between 2.70 and 3.70 million years ago.[38] The tiger's whole genome sequencing shows repeated sequences that parallel those in other cat genomes.[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][37] 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.[37] 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.[37] 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 have a common ancestor that lived between 108,000 and 72,000 years ago.[29] 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 in a 2016 study using 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]

Hybrids

Further information: Felid hybrid and Panthera hybrid

Captive tigers were bred with lions to create hybrids that share physical and behavioural qualities of both parent species; 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.[46] 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, and the lioness passes on a growth inhibiting gene; hence, tigons are around the same size as either species.[47] Since they often develop life threatening birth defects and can easily become obese, breeding these hybrids is regarded as unethical.[46]

Characteristics

Siberian tiger in Aalborg Zoo, Denmark

Bengal tiger skeleton on display at the Museum of Osteology

The tiger has a typical felid morphology. Its body is muscular with shortened legs, strong forelimbs, broad paws, a large head and a tail that is about half the length of the rest of its body.[48][49] It has five digits on the front feet and four on the back, all of which have retractile claws that are compact and curved. The ears are rounded, while the eyes have a round pupil.[48] 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][48] 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).[48][51]

The tiger is the largest living felid species, with the Siberian and Bengal tigers being the largest.[48] 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). Island tigers are the smallest; the Sumatran tiger has 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.[52] The extinct Bali tiger was even smaller.[48] 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.[48][11]

Coat

Tiger coat

The tiger's coat is generally coarse and relatively thin, though the Siberian tiger has a thick winter coat.[48][53] It has a mane-like heavy growth of fur around the neck and jaws and long whiskers, especially in males.[48] It has an orange colouration that varies from yellowish to reddish.[54] White fur covers the ventral surface, along with parts of the face.[48][55] It also has a prominent white spot on the back of the ears which are surrounded by black.[48] The tiger is marked with distinctive black or dark brown stripes, the patterns of which are unique for each individual.[48][56] The stripes are mostly vertical, but those on the limbs and forehead are horizontal. 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 may split up or split and fuse again. Tail stripes are thick bands and a black tip marks the end.[57]

Stripes are advantageous for camouflage in vegetation with vertical patterns of light and shade, such as trees, reeds and tall grass.[56][58] This is supported by a 1987 Fourier analysis study which concluded that the spatial frequencies of tiger stripes line up with their environment.[59] 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 patterns among felids.[60] The orange colour may also aid in concealment, as the tiger's prey are dichromats, and thus they may perceive the cat as green and blended in with the vegetation.[61]

Colour variations

Pseudo-melanistic white tiger

The three colour variants – 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 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 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; 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 habitat

A Bengal tiger in the Sundarbans

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.[48] 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.[52] 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, 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 ecology

Tiger 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 in the early morning with a peak around dawn.[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 exhibited a high temporal overlap with ungulate species.[81]

Tigers groom themselves, maintaining their coats by licking them and spreading oil from their sebaceous glands.[82] It will take to water, particularly on hot days. It is a powerful swimmer and easily transverses rivers as wide as 8 km (5.0 mi).[56] Adults only occasionally climb trees, but have been recorded climbing 10 m (33 ft) up a smooth pipal tree.[48] 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.[83]

Social spacing

Adult tigers lead largely solitary lives and establish and maintain 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.[48][56] Two females in the Sundarbans had home ranges of 10.6 and 14.1 km2 (4.1 and 5.4 sq mi).[84] 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.[85] 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.[86] When tigresses in the same reserve had cubs of up to four months of age, they reduced their home ranges and steadily enlarged them until their offspring were 13–18 months old.[87]

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.[88] Young tigresses establish their first home ranges 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.[89] 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).[90] A young male may have to live as a transient in another male's home range until he is older and strong enough to challenge the resident male. Young males have an annual mortality rate of up to 35 percent. By contrast, young female tigers die at a rate of only around 5 percent.[89] 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.[56][91][92][93] Scent markings also allow an individual to pick up information on another's identity. A tigress in oestrus signals her readiness to mate by scent marking more frequently and increasing her vocalisations. Unclaimed home ranges, particularly those that belonged to a deceased individual, can be taken over in days or weeks.[56]

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 outright violence. Once dominance has been established, a male may tolerate a subordinate within his range, as long as they do not live in close quarters. 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 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]

Communication

Tiger baring teeth as a sign of aggression

Captive 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.[97][48] Both males and females show a flehmen response, a characteristic grimace, when sniffing urine markings. Males also use the flehmen to detect the markings made by tigresses in oestrus.[48] Tigers will move around their ears to display the white spots, particularly during aggressive encounters and between mothers and cubs.[98] 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.[99]

Tigers are normally silent but can produce numerous vocalisations.[100][101] 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.[48][102] Aggressive encounters involve growling, snarling and hissing.[103] An explosive "coughing roar" or "coughing snarl" is emitted through an open mouth and exposed teeth.[48][103][98] In friendlier situations, tigers prusten, a soft, low-frequency snorting sound similar to purring in smaller cats.[104] Tiger mothers communicate with their cubs by grunting, while cubs call back with miaows.[105] When startled they emit a "woof" sound. They produce a deer-like "pok" sound for unknown reasons, but most often at kills.[106][107]

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 ungulates like gaur[108] and opportunistically, smaller prey like monkeys, peafowl and other ground-based birds, porcupines and fish.[48][56] Occasional attacks on Asian elephants and Indian rhinoceros have also been reported.[109] More often, tigers take the more vulnerable small calves.[110] They sometimes prey on livestock and dogs in close proximity to settlements.[48] They occasionally consume vegetation, fruit and minerals for dietary fibre.[111]

Tigers learn to hunt from their mothers, which is important but not necessary for their success.[112] Depending on the prey, they typically kill weekly though mothers must kill more often.[52] They usually hunt alone, but families hunt together when cubs are old enough.[113] They search for prey using vision and hearing.[114] A tiger will also wait at a watering hole for prey to come by, particularly during hot summer days.[115][116] 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,[117] usually within 30 m (98 ft).[52] If the prey spots it before then, the cat does not pursue further.[115] A tiger can sprint 56 km/h (35 mph) and leap 10 m (33 ft);[118][119] it is not a long-distance runner and gives up a chase if prey outpaces it over a certain distance.[115]

Two 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. The tiger generally applies a bite to the throat until its victim dies of strangulation.[48][120][121] It has an average bite force at the canine tips of 1234.3 newtons.[122] Holding onto the throat puts the cat out of reach of horns, antlers, tusks and hooves.[120][123] 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.[124] They kill small prey with a bite to the back of the neck or skull.[125][52] Estimates of the success rate for hunting tigers range from a of low 5% to a high of 50%. They are sometimes killed or injured by large or dangerous prey like gaur, buffalo and boar.[52]

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.[126]

Enemies and competitors

An 1807 illustration of dholes attacking a tiger

Tigers sometimes kill other predators.[127][128] 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.[129] Large dhole packs may kill tigers.[130] Tigers appear to inhabit the deep parts of a forest while these smaller predators are pushed closer to the fringes.[131] The three predators coexist by hunting different prey.[132] 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.[133] Leopards can live successfully in tiger habitat when there is abundant food and vegetation cover.[132][134] Nevertheless, leopards avoid areas where tigers roam and are less common where tigers are numerous.[127][135][136]

Tigers tend to be wary of sloth bears, with their sharp claws, quickness and ability to stand on two legs. Tigers sometimes prey on sloth bears by ambushing them when they are feeding at termite mounds.[137] Siberian tigers attack, kill and prey on Ussuri brown and black bears.[48] Brown bears frequently track down tigers to usurp their kills, with occasional fatal outcomes for the tiger.[138][139]

Reproduction and life cycle

"Tiger cub" redirects here. For other uses, see Tiger Cub.

A 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, in between three and nine week intervals.[48] A resident male mates with all the females within his home range, who signal their receptiveness by roaring and marking.[140][141] Younger, transient males are also attracted, leading to a fight in which the more dominant male drives the usurper off.[142][140] 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 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.[140] Tiger pairs may stay together for up to four days and mate multiple times.[143] Gestation lasts around or over three months.[48]

A tigress gives birth in a secluded location, be it in dense vegetation, in a cave or under a rocky shelter.[144] Litters consist of as many as seven cubs, but two or three are more typical.[142][144] Newborn cubs weigh 785–1,610 g (27.7–56.8 oz), and are blind and altricial.[144] The mother licks and cleans her cubs, suckles them and viciously defends them from any potential threat.[142] Young open their eyes in a week, can leave the denning site in two months and around the same time they start eating meat.[142][145] The mother only leaves them alone to hunt, and even then she does not travel far.[146] 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. Cubs may die at a rate of 50% during these early months, sources of morality include predators, floods and fires.[147][148]

A 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, and she will guide them to the kill. Cubs 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.[149] 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.[147] A cub can make a kill as early as 11 months and reach independence around 18 to 24 months of age; males become independent earlier than females.[150] Radio-collared tigers in Chitwan started dispersing from their natal areas at the age of 19 months.[90] Young females are sexually mature at three to four years, whereas males are at four to five years.[48] Generation length of the tiger is about 7–10 years.[151] Wild Bengal tigers live 12–15 years.[152] Data from the International Tiger Studbook 1938–2018 indicate that captive tigers lived up to 19 years.[153]

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.[154][155] One male was recorded looking after orphaned cubs whose mother had died.[156] By defending his home range, the male protects the females and cubs from other males.[157] When a new male takes over, dependent cubs are at risk of being killed, as the male attempts to sire his own young with the females. Older female cubs are tolerated but males are treated as potential competitors.[158]

Health and diseases

Tigers are recorded as hosts for various parasites including Diphyllobothrium erinacei, Taenia pisiformis and Toxocara species in India and Physaloptera preputialis, Dirofilaria ursi and Uiteinarta species in Siberia.[48] Canine distemper is known to occur in Siberian tigers.[159] 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.[160] Blood samples from 11 adult tigers in Nepal showed antibodies for canine parvovirus-2, feline herpesvirus, feline coronavirus, leptospirosis and Toxoplasma gondii.[161]

Threats

The tiger has been listed as Endangered on the IUCN Red List since 1986, as 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.[162] In the Tanintharyi Region of southern Myanmar, deforestation coupled with mining activities and high hunting pressure threatens the tiger population in the area.[163] 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 have no longer harboured tigers since at least 2013.[164] 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.[165] 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.[166] 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.[167] 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.[168][169]

A Javan tiger skin on display

Tiger populations in India have been targeted by poachers since the 1990s and were extirpated in two tiger reserves in 2005 and 2009.[170] 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.[171] 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.[172] 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.[173] 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 vital for prey species to survive in winter.[174]

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.[175] 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.[176] Seizures in Nepal between January 2011 and December 2015 included 585 pieces of tiger body parts and two whole bodies in 19 districts.[177] 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.[178] 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.[179]

Demand for tiger parts for use in traditional Chinese medicine has also been cited as a major threat to tiger populations.[180] 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.[181] 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.[182] Tiger bone glue is the prevailing tiger product purchased for medicinal purposes in Hanoi and Ho Chi Minh City.[183] "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.[184]

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.[185][186][187][188][189]

Conservation

Relationship with humans