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The cheetah (Acinonyx jubatus) is a large cat and the fastest land animal. It has a tawny to creamy white or pale buff fur that is marked with evenly spaced, solid black spots. The head is small and rounded, with a short snout and black tear-like facial streaks. It reaches Template:Cvt at the shoulder, and the head-and-body length is between Template:Cvt. Adults weigh between Template:Cvt. The cheetah is capable of running at Template:Cvt; it has evolved specialized adaptations for speed, including a light build, long thin legs and a long tail.
The cheetah was first described in the late 18th century. Four subspecies are recognised today that are native to Africa and central Iran. An African subspecies was introduced to India in 2022. It is now distributed mainly in small, fragmented populations in northwestern, eastern and southern Africa and central Iran. It lives in a variety of habitats such as savannahs in the Serengeti, arid mountain ranges in the Sahara, and hilly desert terrain.
The cheetah lives in three main social groups: females and their cubs, male "coalitions", and solitary males. While females lead a nomadic life searching for prey in large home ranges, males are more sedentary and instead establish much smaller territories in areas with plentiful prey and access to females. The cheetah is active during the day, with peaks during dawn and dusk. It feeds on small- to medium-sized prey, mostly weighing under Template:Cvt, and prefers medium-sized ungulates such as impala, springbok and Thomson's gazelles. The cheetah typically stalks its prey within Template:Cvt before charging towards it, trips it during the chase and bites its throat to suffocate it to death. It breeds throughout the year. After a gestation of nearly three months, females give birth to a litter of three or four cubs. Cheetah cubs are highly vulnerable to predation by other large carnivores. They are weaned at around four months and are independent by around 20 months of age.
The cheetah is threatened by habitat loss, conflict with humans, poaching and high susceptibility to diseases. The global cheetah population was estimated at 6,517 individuals in 2021; it is listed as Vulnerable on the IUCN Red List. It has been widely depicted in art, literature, advertising, and animation. It was tamed in ancient Egypt and trained for hunting ungulates in the Arabian Peninsula and India. It has been kept in zoos since the early 19th century.
EtymologyEdit
The vernacular name "cheetah" is derived from Hindustani Template:Langx and Template:Langx ({{#invoke:Lang|lang}}).<ref>Template:Cite book</ref> This in turn comes from Template:Langx ({{#invoke:Lang|lang}}) meaning 'variegated', 'adorned' or 'painted'.<ref>Template:Cite book</ref> In the past, the cheetah was often called "hunting leopard" because they could be tamed and used for coursing.<ref name=marker1>Template:Cite book</ref> The generic name Acinonyx probably derives from the combination of two Greek words: {{#invoke:Lang|lang}} ({{#invoke:Lang|lang}}) meaning 'unmoved' or 'motionless', and {{#invoke:Lang|lang}} ({{#invoke:Lang|lang}}) meaning 'nail' or 'hoof'.<ref>Template:Cite book</ref> A rough translation is "immobile nails", a reference to the cheetah's limited ability to retract its claws.<ref name=Rosevear1974>Template:Cite book</ref> A similar meaning can be obtained by the combination of the Greek prefix a– (implying a lack of) and {{#invoke:Lang|lang}} ({{#invoke:Lang|lang}}) meaning 'to move' or 'to set in motion'.<ref name="skinner">Template:Cite book</ref> The specific name {{#invoke:Lang|lang}} is Latin for 'crested, having a mane'.<ref>Template:Cite book</ref>
A few old generic names such as Cynailurus and Cynofelis allude to the similarities between the cheetah and canids.<ref name="marker7">Template:Cite book</ref>
TaxonomyEdit
In 1777, Johann Christian Daniel von Schreber described the cheetah based on a skin from the Cape of Good Hope and gave it the scientific name Felis jubatus.<ref>Template:Cite book</ref> Joshua Brookes proposed the generic name Acinonyx in 1828.<ref>Template:Cite book</ref> In 1917, Reginald Innes Pocock placed the cheetah in a subfamily of its own, Acinonychinae,<ref name=Pocock1917>Template:Cite journal</ref> given its striking morphological resemblance to the greyhound and significant deviation from typical felid features; the cheetah was classified in Felinae in later taxonomic revisions.<ref name=caro1994/>
In the 19th and 20th centuries, several cheetah zoological specimens were described; some were proposed as subspecies.<ref name=MSW3>Template:MSW3 Wozencraft</ref> A South African specimen with notably dense fur was proposed as (Felis lanea) by Philip Sclater in 1877 and became known as the "woolly cheetah".<ref>Template:Cite journal</ref> Its classification as a species was mostly disputed.<ref>Template:Cite book</ref> There has been considerable confusion in the nomenclature of the cheetah and leopard (Panthera pardus) as authors often confused the two; some considered "hunting leopards" an independent species, or equal to the leopard.<ref>Template:Cite book</ref><ref>Template:Cite book</ref>
SubspeciesEdit
In 1975, five cheetah subspecies were considered valid taxa: A. j. hecki, A. j. jubatus, A. j. raineyi, A. j. soemmeringii and A. j. venaticus.<ref name=Catsg2017/> In 2011, a phylogeographic study found minimal genetic variation between A. j. jubatus and A. j. raineyi; only four subspecies were identified.<ref name=subspecies>Template:Cite journal</ref> In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group revised felid taxonomy and recognised these four subspecies as valid. Their details are tabulated below:<ref name=Catsg2017/>
| Subspecies | Details | Image | Range map | |
|---|---|---|---|---|
| Southeast African cheetah (A. j. jubatus) Template:Small syn. A. j. raineyi Template:Small<ref name=heller>Template:Cite journal</ref> |
The nominate subspecies;<ref name=MSW3/> it genetically diverged from the Asiatic cheetah 67,000–32,000 years ago.<ref name="subspecies"/> As of 2016, the largest population of nearly 4,000 individuals is sparsely distributed in Angola, Botswana, Mozambique, Namibia, South Africa and Zambia.<ref name=durant2017/> | File:Cheetah, Masai Mara (52448476886).jpg | File:Acinonyx jubatus jubatus range IUCN 2015.png | |
| Asiatic cheetah (A. j. venaticus) Template:Small<ref name=griffith>Template:Cite book</ref> |
This subspecies is confined to central Iran, and is the only surviving cheetah population in Asia.<ref name=marker4>Template:Cite book</ref> As of 2022, only 12 individuals were estimated to survive in Iran, nine of which are males and three of which are females.<ref>{{#invoke:citation/CS1|citation | CitationClass=web
}}</ref> |
File:Iranian Cheetah roars.jpg | File:Acinonyx jubatus venaticus range IUCN 2015.png |
| Northeast African cheetah (A. j. soemmeringii) Template:Small<ref>Template:Cite book</ref> |
This subspecies occurs in the northern Central African Republic, Chad, Ethiopia and South Sudan in small and heavily fragmented populations; in 2016, the largest population of 238 individuals occurred in the northern CAR and southeastern Chad. It diverged genetically from the southeast African cheetah 72,000–16,000 years ago.<ref name=subspecies/> | File:Cheetah in the shade DVIDS147321.jpg | File:Acinonyx jubatus soemmeringii range IUCN 2015.png | |
| Northwest African cheetah (A. j. hecki) Template:Small<ref>Template:Cite book</ref> |
This subspecies occurs in Algeria, Benin, Burkina Faso, Mali and Niger.<ref name=iucn/> In 2016, the largest population of 191 individuals occurred in Adrar des Ifoghas, Ahaggar and Tassili n'Ajjer in south-central Algeria and northeastern Mali.<ref name=marker4/> It is listed as critically endangered on the IUCN Red List.<ref name=iucn3>Template:Cite iucn</ref> | File:Acinonyx jubatus ssp. hecki in Idlès 1.jpg | File:Acinonyx jubatus hecki range IUCN 2015.png |
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Phylogeny and evolutionEdit
The cheetah's closest relatives are the cougar (Puma concolor) and the jaguarundi (Herpailurus yagouaroundi).<ref name="Catsg2017">Template:Cite journal</ref> Together, these three species form the Puma lineage, one of the eight lineages of the extant felids; the Puma lineage genetically diverged from the rest 6.7 mya. The sister group of the Puma lineage is a clade of smaller Old World cats that includes the genera Felis, Otocolobus and Prionailurus.<ref name="bcw2">Template:Cite book</ref>
The oldest cheetah fossils, excavated in eastern and southern Africa, date to 3.5–3 mya; the earliest known specimen from South Africa is from the lowermost deposits of the Silberberg Grotto (Sterkfontein).<ref name=mammal/><ref name=skinner/> Though incomplete, these fossils indicate forms larger but less cursorial than the modern cheetah.<ref name="marker2">Template:Cite book</ref> The first occurrence of the modern species A. jubatus in Africa may come from Cooper's D, a site in South Africa dating back to 1.5 to 1.4 Ma, during the Calabrian stage.<ref>Template:Cite journal</ref> Fossil remains from Europe are limited to a few Middle Pleistocene specimens from Hundsheim (Austria) and Mosbach Sands (Germany).<ref name="hemmer">Template:Cite journal</ref> Cheetah-like cats are known from as late as 10,000 years ago from the Old World. The giant cheetah (A. pardinensis), significantly larger and slower compared to the modern cheetah, occurred in Eurasia and eastern and southern Africa in the Villafranchian period roughly 3.8–1.9 mya.<ref name=caro1994/><ref>Template:Cite journal</ref> In the Middle Pleistocene a smaller cheetah, A. intermedius, ranged from Europe to China.<ref name=mammal>Template:Cite journal</ref> The modern cheetah appeared in Africa around 1.9 mya; its fossil record is restricted to Africa.<ref name=marker2/>
Extinct North American cheetah-like cats had historically been classified in Felis, Puma or Acinonyx; two such species, F. studeri and F. trumani, were considered to be closer to the puma than the cheetah, despite their close similarities to the latter. Noting this, palaeontologist Daniel Adams proposed Miracinonyx, a new subgenus under Acinonyx, in 1979 for the North American cheetah-like cats;<ref name="adams"/> this was later elevated to genus rank.<ref name="Valkenburgh1990" /> Adams pointed out that North American and Old World cheetah-like cats may have had a common ancestor, and Acinonyx might have originated in North America instead of Eurasia.<ref name="adams">Template:Cite journal</ref> However, subsequent research has shown that Miracinonyx is phylogenetically closer to the cougar than the cheetah;<ref name=sabre>Template:Cite journal</ref> the similarities to cheetahs have been attributed to parallel evolution.<ref name="bcw2"/>
The three species of the Puma lineage may have had a common ancestor during the Miocene (roughly 8.25 mya).<ref name=adams/><ref name=johnson>Template:Cite journal</ref> Some suggest that North American cheetahs possibly migrated to Asia via the Bering Strait, then dispersed southward to Africa through Eurasia at least 100,000 years ago;<ref>Template:Cite journal</ref><ref name="dobrynin">Template:Cite journal</ref><ref>Template:Cite journal</ref> some authors have expressed doubt over the occurrence of cheetah-like cats in North America, and instead suppose the modern cheetah to have evolved from Asian populations that eventually spread to Africa.<ref name=sabre/><ref>Template:Cite journal</ref> The cheetah is thought to have experienced two population bottlenecks that greatly decreased the genetic variability in populations; one occurred about 100,000 years ago that has been correlated to migration from North America to Asia, and the second 10,000–12,000 years ago in Africa, possibly as part of the Late Pleistocene extinction event.<ref name=dobrynin/><ref name="o'brien1987">Template:Cite journal</ref><ref name="o'brien1993">Template:Cite journal</ref>
GeneticsEdit
The diploid number of chromosomes in the cheetah is 38, the same as in most other felids.<ref name=Geptner1972/> The cheetah was the first felid observed to have unusually low genetic variability among individuals,<ref name="bcw4">Template:Cite book</ref> which has led to poor breeding in captivity, increased spermatozoal defects, high juvenile mortality and increased susceptibility to diseases and infections.<ref name="o'brien1985">Template:Cite journal</ref><ref name=obrien2017>Template:Cite journal</ref> A prominent instance was the deadly feline coronavirus outbreak in the cheetah breeding facility at Wildlife Safari in Winston, Oregon in 1983 which had a mortality rate of 60%, higher than that recorded for previous epizootics of feline infectious peritonitis in any felid.<ref>Template:Cite journal</ref> The remarkable homogeneity in cheetah genes has been demonstrated by experiments involving the major histocompatibility complex (MHC); unless the MHC genes are highly homogeneous in a population, skin grafts exchanged between a pair of unrelated individuals would be rejected. Skin grafts exchanged between unrelated cheetahs are accepted well and heal, as if their genetic makeup were the same.<ref name=yuhki>Template:Cite journal</ref><ref name=tears>Template:Cite book</ref>
The low genetic diversity is thought to have been created by two population bottlenecks from about 100,000 years and about 12,000 years ago, respectively.<ref name="dobrynin" /> The resultant level of genetic variation is around 0.1–4% of average living species, lower than that of Tasmanian devils, Virunga gorillas, Amur tigers, and even highly inbred domestic cats and dogs.<ref name=obrien2017/>
Selective retention of gene variants (duplication) has been found in 10 gene candidates to explain energetics and anabolism related to muscle specialization in cheetahs:
- Regulation of muscle contraction (Five genes: ADORA1, ADRA1B, CACNA1C, RGS2, SCN5A).
- Physiological stress response (Two genes: ADORA1, TAOK2).
- Negative regulation of catabolic process (Four genes: APOC3, SUFU, DDIT4, PPARA).<ref name="dobrynin" /><ref name="obrien2017" />
Potentially harmful mutations has been found in a gene related to spermatogenesis (AKAP4). This could explain the high proportion of abnormal sperma in male cheetahs and poor reproductive success in the species.<ref name="dobrynin" /><ref name="obrien2017" />
King cheetahEdit
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The king cheetah is a variety of cheetah with a rare mutation for cream-coloured fur marked with large, blotchy spots and three dark, wide stripes extending from the neck to the tail.<ref name=thompson>Template:Cite book</ref> In Manicaland, Zimbabwe, it was known as nsuifisi and thought to be a cross between a leopard and a hyena.<ref name=bottriell/> In 1926, Major A. Cooper wrote about a cheetah-like animal he had shot near modern-day Harare, with fur as thick as that of a snow leopard and spots that merged to form stripes. He suggested it could be a cross between a leopard and a cheetah. As more such individuals were observed it was seen that they had non-retractable claws like the cheetah.<ref name=pocock/><ref name=heuvelmans>Template:Cite book</ref>
In 1927, Pocock described these individuals as a new species by the name of Acinonyx rex ("king cheetah").<ref name="pocock">Template:Cite journal</ref> However, in the absence of proof to support his claim, he withdrew his proposal in 1939. Abel Chapman considered it a colour morph of the normally spotted cheetah.<ref name=catsg>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Since 1927, the king cheetah has been reported five more times in the wild in Zimbabwe, Botswana and northern Transvaal; one was photographed in 1975.<ref name="bottriell">Template:Cite book</ref>
In 1981, two female cheetahs that had mated with a wild male from Transvaal at the De Wildt Cheetah and Wildlife Centre (South Africa) gave birth to one king cheetah each; subsequently, more king cheetahs were born at the centre.<ref name = catsg/> In 2012, the cause of this coat pattern was found to be a mutation in the gene for transmembrane aminopeptidase (Taqpep), the same gene responsible for the striped "mackerel" versus blotchy "classic" pattern seen in tabby cats.<ref>Template:Cite journal</ref> The appearance is caused by reinforcement of a recessive allele; hence if two mating cheetahs are heterozygous carriers of the mutated allele, a quarter of their offspring can be expected to be king cheetahs.<ref name="wcw">Template:Cite book</ref>
CharacteristicsEdit
The cheetah is a lightly built, spotted cat characterised by a small rounded head, a short snout, black tear-like facial streaks, a deep chest, long thin legs and a long tail. Its slender, canine-like form is highly adapted for speed, and contrasts sharply with the robust build of the genus Panthera.<ref name=marker7/><ref name="bcw3">Template:Cite book</ref> Cheetahs typically reach Template:Cvt at the shoulder and the head-and-body length is between Template:Cvt.<ref name=skinner/><ref name=kingdon/><ref name=nowak>Template:Cite book</ref> The weight can vary with age, health, location, sex and subspecies; adults typically range between Template:Cvt.<ref name="marker2003" /><ref name="Schaller-1972" /> Cubs born in the wild weigh Template:Cvt at birth, while those born in captivity tend to be larger and weigh around Template:Cvt.<ref name=marker7/><ref name=kingdon/> The cheetah is sexually dimorphic, with males larger and heavier than females, but not to the extent seen in other large cats; females have a much lower body mass index than males.<ref name="marker2003">Template:Cite journal</ref> Studies differ significantly on morphological variations among the subspecies.<ref name=marker2003/>
The coat is typically tawny to creamy white or pale buff (darker in the mid-back portion).<ref name=skinner/><ref name=kingdon/> The chin, throat and underparts of the legs and the belly are white and devoid of markings. The rest of the body is covered with around 2,000 evenly spaced, oval or round solid black spots, each measuring roughly Template:Cvt.<ref name=wcw/><ref name=hunterwcw/> Each cheetah has a distinct pattern of spots which can be used to identify unique individuals.<ref name=nowak/> Besides the clearly visible spots, there are other faint, irregular black marks on the coat.<ref name=hunterwcw/> Newly born cubs are covered in fur with an unclear pattern of spots that gives them a dark appearance—pale white above and nearly black on the underside.<ref name=marker7/> The hair is mostly short and often coarse, but the chest and the belly are covered in soft fur; the fur of king cheetahs has been reported to be silky.<ref name=skinner/><ref name=Estes>Template:Cite book</ref> There is a short, rough mane, covering at least Template:Cvt along the neck and the shoulders; this feature is more prominent in males. The mane starts out as a cape of long, loose blue to grey hair in juveniles.<ref name=wcw/><ref name=Estes/> Melanistic cheetahs are rare and have been seen in Zambia and Zimbabwe.<ref name=hunterwcw/> In 1877–1878, Sclater described two partially albino specimens from South Africa.<ref name=wcw/>
The head is small and more rounded compared to other big cats.<ref name="mills">Template:Cite book</ref> Saharan cheetahs have canine-like slim faces.<ref name="hunterwcw">Template:Cite book</ref> The ears are small, short and rounded; they are tawny at the base and on the edges and marked with black patches on the back. The eyes are set high and have round pupils.<ref name=nowak/> The whiskers, shorter and fewer than those of other felids, are fine and inconspicuous.<ref name=Montgomery>Template:Cite book</ref> The pronounced tear streaks (or malar stripes), unique to the cheetah, originate from the corners of the eyes and run down the nose to the mouth. The role of these streaks is not well understood—they may protect the eyes from the sun's glare (a helpful feature as the cheetah hunts mainly during the day), or they could be used to define facial expressions.<ref name=hunterwcw/> The exceptionally long and muscular tail, with a bushy white tuft at the end, measures Template:Cvt.<ref name = Stuart/> While the first two-thirds of the tail are covered in spots, the final third is marked with four to six dark rings or stripes.<ref name=wcw/><ref name="arnold">Template:Cite book</ref>
The cheetah is superficially similar to the leopard, which has a larger head, fully retractable claws, rosettes instead of spots, lacks tear streaks and is more muscular.<ref name="hilde">Template:Cite book</ref><ref name="foley">Template:Cite book</ref> Moreover, the cheetah is taller than the leopard. The serval also resembles the cheetah in physical build, but is significantly smaller, has a shorter tail and its spots fuse to form stripes on the back.<ref>Template:Cite book</ref> The cheetah appears to have evolved convergently with canids in morphology and behaviour; it has canine-like features such as a relatively long snout, long legs, a deep chest, tough paw pads and blunt, semi-retractable claws.<ref>Template:Cite book</ref><ref>Template:Cite journal</ref> The cheetah has often been likened to the greyhound, as both have similar morphology and the ability to reach tremendous speeds in a shorter time than other mammals,<ref name="Estes" /><ref name=Stuart>Template:Cite book</ref> but the cheetah can attain much higher maximum speeds.<ref>Template:Cite journal</ref>
Internal anatomyEdit
Template:Multiple image Sharply contrasting with the other big cats in its morphology, the cheetah shows several specialized adaptations for prolonged chases to catch prey at some of the fastest speeds reached by land animals.<ref name="claw">Template:Cite journal</ref> Its light, streamlined body makes it well-suited to short, explosive bursts of speed, rapid acceleration, and an ability to execute extreme changes in direction while moving at high speed.<ref name=cheathsr>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref name="WilsonBiologyLetters">Template:Cite journal</ref> The large nasal passages, accommodated well due to the smaller size of the canine teeth, ensure fast flow of sufficient air, and the enlarged heart and lungs allow the enrichment of blood with oxygen in a short time. This allows cheetahs to rapidly regain their stamina after a chase.<ref name=mammal/> During a typical chase, their respiratory rate increases from 60 to 150 breaths per minute.<ref name="O'Brien">Template:Cite journal</ref> The cheetah has a fast heart rate, averaging 126–173 beats per minute at resting without arrhythmia.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Moreover, the reduced viscosity of the blood at higher temperatures (common in frequently moving muscles) could ease blood flow and increase oxygen transport.<ref>Template:Cite journal</ref> While running, in addition to having good traction due to their semi-retractable claws, cheetahs use their tail as a rudder-like means of steering that enables them to make sharp turns, necessary to outflank antelopes which often change direction to escape during a chase.<ref name=wcw/><ref name=mills/> The protracted claws increase grip over the ground, while rough paw pads make the sprint more convenient over tough ground. The limbs of the cheetah are longer than what is typical for other cats its size; the thigh muscles are large, and the tibia and fibula are held close together making the lower legs less likely to rotate. This reduces the risk of losing balance during runs, but compromises the cat's ability to climb trees. The highly reduced clavicle is connected through ligaments to the scapula, whose pendulum-like motion increases the stride length and assists in shock absorption. The extension of the vertebral column can add as much as Template:Cvt to the stride length.<ref name=hildebrand>Template:Cite journal</ref><ref name=bertram>Template:Cite journal</ref>
Muscle tissue has been analyzed in the cheetah and it has been found that there are little differences in type II muscle fiber concentration, anaerobic lactate dehydrogenase enzyme activity, and glycogen concentration between sexes.<ref name=":0">Template:Cite journal</ref>
The cheetah resembles the smaller cats in cranial features, and in having a long and flexible spine, as opposed to the stiff and short one in other large felids.<ref name=mammal/> The roughly triangular skull has light, narrow bones and the sagittal crest is poorly developed, possibly to reduce weight and enhance speed. The mouth can not be opened as widely as in other cats given the shorter length of muscles between the jaw and the skull.<ref name=wcw/><ref name="hilde"/> A study suggested that the limited retraction of the cheetah's claws may result from the earlier truncation of the development of the middle phalanx bone in cheetahs.<ref name=claw/>
The cheetah has a total of 30 teeth; the dental formula is Template:DentalFormula. The small, flat canines are used to bite the throat and suffocate the prey. A study gave the bite force quotient (BFQ) of the cheetah as 119, close to that for the lion (112), suggesting that adaptations for a lighter skull may not have reduced the power of the cheetah's bite.<ref name=mammal/><ref name=marker7/> Unlike other cats, the cheetah's canines have no gap or diastema behind them when the jaws close, as the top and bottom cheek teeth show extensive overlap.<ref name="bcw3" /> Cheetahs have relatively elongated, blade-like shape carnassial teeth, with reduced lingual cusps; this may have been an adaptation to consume quickly the flesh of a prey before more heavy-built predators from other species arrive to take it from them.<ref>Template:Cite book</ref> The slightly curved claws, shorter and straighter than those of other cats, lack a protective sheath and are partly retractable.<ref name=wcw/><ref name=nowak/> The claws are blunt due to lack of protection,<ref name=hunterwcw/> but the large and strongly curved dewclaw is remarkably sharp.<ref name=dewclaw>Template:Cite journal</ref> Cheetahs have a high concentration of nerve cells arranged in a band in the centre of the eyes, a visual streak, the most efficient among felids. This significantly sharpens the vision and enables the cheetah to swiftly locate prey against the horizon.<ref name=bcw3/><ref>Template:Cite journal</ref> The cheetah is unable to roar due to the presence of a sharp-edged vocal fold within the larynx.<ref name=mammal/><ref name=hast>Template:Cite journal</ref>
In stressful situations, the cheetah has a lower cortisol level than the leopard, indicating better stress response; it also has lower immunoglobulin G and Serum amyloid A levels but a higher lysozyme level and a higher bacterial killing capacity than the leopard, indicating a poorer adaptive and induced innate immune systems but a better constitutive innate immune system; its constitutive innate immune system compensates for its low variation of major histocompatibility complex and poorer immune adaptability.<ref>Template:Cite journal</ref>
Speed and accelerationEdit
The cheetah is the world's fastest land animal.<ref name=gonyea>Template:Cite journal</ref><ref name=hudson>Template:Cite journal</ref> Estimates of the maximum speed attained range from Template:Cvt.<ref name=wcw/><ref name=nowak/> A commonly quoted value is Template:Cvt, recorded in 1957, but this measurement is disputed.<ref>Template:Cite newsTemplate:Cbignore</ref> In 2012, an 11-year-old cheetah from the Cincinnati Zoo set a world record by running Template:Cvt in 5.95 seconds over a set run, recording a maximum speed of Template:Cvt.<ref name=Pappas>Template:Cite news</ref>
Cheetahs equipped with GPS collars hunted at speeds during most of the chase much lower than the highest recorded speed; their run was interspersed with a few short bursts of a few seconds when they attained peak speeds. The average speed recorded during the high speed phase was Template:Cvt, or within the range Template:Cvt including error. The highest recorded value was Template:Cvt. A hunt consists of two phases, an initial fast acceleration phase when the cheetah tries to catch up with the prey, followed by slowing down as it closes in on it, the deceleration varying by the prey in question. The initial linear acceleration observed was 13 m/s², more than twice than 6 m/s² of horses and greater than 10 m/s² of greyhounds.<ref name=":1" /><ref>Template:Cite journal</ref> Cheetahs can increase up 3 m/s (10.8 km/h) and decrease up 4 m/s (14.4 km/h) in a single stride.<ref name="Wilson_al2013">Template:Cite journal</ref> Speed and acceleration values for a hunting cheetah may be different from those for a non-hunter because while engaged in the chase, the cheetah is more likely to be twisting and turning and may be running through vegetation.<ref name=Wilson_al2013/><ref name="WilsonBiologyLetters" /> The speeds of more than 100 km/h attained by the cheetah may be only slightly greater than those achieved by the pronghorn at Template:Cvt<ref>Template:Cite book</ref> and the springbok at Template:Cvt,<ref>Template:Cite book</ref> but the cheetah additionally has an exceptional acceleration, can go from 0–97 km/h (0–60 mph) in less than 3 seconds, "faster than a Ferrari".<ref name="Philips, J. A.-1997">Template:Cite journal</ref><ref>Template:Cite journal</ref> For comparison, polo horses can go from 0 to 36 km/h in 3.6 seconds.<ref>Template:Cite journal</ref>
One stride of a galloping cheetah measures Template:Cvt; the stride length and the number of jumps increases with speed.<ref name=wcw/> During more than half the duration of the sprint, the cheetah has all four limbs in the air, increasing the stride length.<ref name=taylor>Template:Cite book</ref> Running cheetahs can retain up to 90% of the heat generated during the chase. A 1973 study suggested the length of the sprint is limited by excessive build-up of body heat when the body temperature reaches Template:Cvt. However, a 2013 study recorded the average temperature of cheetahs after hunts to be Template:Cvt, suggesting high temperatures need not cause hunts to be abandoned.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>
The running speed of Template:Cvt of the cheetah was obtained as an result of a single run of one individual by dividing the distance traveled for time spent. The run lasted 2.25 seconds and was supposed to have been Template:Cvt long, but was later found to have been Template:Cvt long. It was therefore discredited for a faulty method of measurement.<ref name=Hildebrand1959>Template:Cite journal</ref> Cheetahs have subsequently been measured at running at a speed of Template:Cvt as the fastest speed from three runs including in opposite direction, for a single individual, over a marked Template:Cvt course, even starting the run Template:Cvt behind the start line, starting the run already running on the course. Again dividing the distance by time, but this time to determine the maximum sustained speed, completing the runs in an time of 7.0, 6.9 and 7.2 seconds. Being a more accurate method of measurement, this test was made in 1965 but published in 1997.<ref name="Sharp-1997">Template:Cite journal</ref>
In 2010, the running speed of 15 cheetahs was measured by means of high speed camera stationed on a tripod and placed at specific points on a track; the cheetahs were chasing a lure and there were several attempts per individual, and their length from the nose to base of the tail was used as a scale. The speed was estimated from the time the tip of the nose appeared until it was no longer visible on camera. The maximum speed recorded was 100.1 km/h for one individual.<ref>Template:Cite journal</ref> Subsequently, with GPS-IMU collars, in 2011 and 2012, running speed was measured for wild cheetahs during hunts with turns and manoeuvres, and the maximum speed recorded was Template:Cvt sustained for 1–2 seconds. The speed was obtained by dividing the length by the time between footfalls of a stride.<ref name=Wilson_al2013/> There are indirect ways to measure how fast a cheetah can run. One case is known of a cheetah that overtook a young male pronghorn. Cheetahs can overtake a running antelope with a Template:Cvt head start. Both animals were clocked at Template:Cvt by speedometer reading while running alongside a vehicle at full speed.<ref name=Hildebrand1959/> Cheetahs can easily capture gazelles galloping at full speed (Template:Cvt).<ref name=Schaller-1972/>
The physiological reasons for speed in cheetahs are:
- Small head and long lumbar region of the spine, 36.8% of the presacral vertebral column.<ref name=marker2/><ref name=Gonyea1976/><ref name=Valkenburgh1990/>
- A tibia and radius longer than the femur and humerus, with a femorotibial index of 101.9–105 and a humeroradial index of 100.1–103.3.<ref name=marker2 /><ref name=Valkenburgh1990>Template:Cite journal</ref><ref name=Gonyea1976>Template:Cite journal</ref>
- Elongated and slender long bones of the limbs, especially femur, tibia, humerus, radius and pelvis, specially the ischium.<ref name=marker7/><ref name=Gonyea1976/><ref name=hudson/>
- Enlarged respiratory passages and frontal sinuses that allow to cool inhaled and exhaled air with each breath, which helps dissipate body heat.<ref>Template:Cite journal</ref>
- A higher concentration of glycolytic fast twitch muscle fibers (Type IIx) than other cats and animals in general.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> A very high LDH activity is indicative of this principally anaerobic muscle metabolism.<ref name="Philips, J. A.-1997" />
- Most of the locomotor muscle mass is concentrated proximally close to the body in shoulders, thighs and spine, and is reduced in shins and forearms. Long tendons finish off the distal locomotor muscles.<ref name=marker7/><ref name=hudson/>
- Muscular hindlimbs form 19.8% of the body mass, whereas the forelimbs form 15.1%.<ref name=hudson/><ref>Template:Cite journal</ref> The hamstrings, quadriceps, adductor muscles of the hip and psoas major muscles are especially large.<ref name="McNeill">Template:Cite book</ref>
- Enlarged Betz cells in the motor cortex M1 and innervating muscle fibers, to fit its predominant type IIx fibers and powerful muscles.<ref>Template:Cite journal</ref>
Ecology and behaviourEdit
Cheetahs are active mainly during the day,<ref name=Estes/> whereas other carnivores such as leopards and lions are active mainly at night;<ref name=hilde/><ref name=Schaller1972>Template:Cite book</ref> These larger carnivores can kill cheetahs and steal their kills;<ref name=wcw/> hence, the diurnal tendency of cheetahs helps them avoid larger predators in areas where they are sympatric, such as the Okavango Delta. In areas where the cheetah is the major predator (such as farmlands in Botswana and Namibia), activity tends to increase at night. This may also happen in highly arid regions such as the Sahara, where daytime temperatures can reach Template:Convert. The lunar cycle can also influence the cheetah's routine—activity might increase on moonlit nights as prey can be sighted easily, though this comes with the danger of encountering larger predators.<ref name=wcw/><ref name=marker8/> Hunting is the major activity throughout the day, with peaks during dawn and dusk.<ref name=hunterwcw/> Groups rest in grassy clearings after dusk. Cheetahs often inspect their vicinity at observation points such as elevations to check for prey or larger carnivores; even while resting, they take turns at keeping a lookout.<ref name=wcw/>
Social organisationEdit
Cheetahs have a flexible and complex social structure and tend to be more gregarious than several other cats (except the lion). Individuals typically avoid one another but are generally amicable; males may fight over territories or access to females in oestrus, and on rare occasions such fights can result in severe injury and death. Females are not social and have minimal interaction with other individuals, barring the interaction with males when they enter their territories or during the mating season. Some females, generally mother and offspring or siblings, may rest beside one another during the day. Females tend to lead a solitary life or live with offspring in undefended home ranges; young females often stay close to their mothers for life but young males leave their mother's range to live elsewhere.<ref name=wcw/><ref name=nowak/><ref name=hunterwcw/>
Some males are territorial, and group together for life, forming coalitions that collectively defend a territory which ensures maximum access to females—this is unlike the behaviour of the male lion who mates with a particular group (pride) of females. In most cases, a coalition will consist of brothers born in the same litter who stayed together after weaning, but biologically unrelated males are often allowed into the group; in the Serengeti, 30% of members in coalitions are unrelated males.<ref name=hunterwcw/> If a cub is the only male in a litter, he will typically join an existing group, or form a small group of solitary males with two or three other lone males who may or may not be territorial. In the Kalahari Desert around 40% of the males live in solitude.<ref name=nowak/><ref name = hunterwcw/>
Males in a coalition are affectionate toward each other, grooming mutually and calling out if any member is lost; unrelated males may face some aversion in their initial days in the group. All males in the coalition typically have equal access to kills when the group hunts together, and possibly also to females who may enter their territory.<ref name=marker9>Template:Cite book</ref> A coalition generally has a greater chance of encountering and acquiring females for mating; however, its large membership demands greater resources than do solitary males.<ref name=nowak/><ref name=hunterwcw/> A 1987 study showed that solitary and grouped males have a nearly equal chance of coming across females, but the males in coalitions are notably healthier and have better chances of survival than their solitary counterparts.<ref name=caro1987>Template:Cite journal</ref>
Male cheetahs seem to be more tolerant to cubs that are not their offspring than other felids, and supposed evidence of infanticide is considered circumstantial.<ref>Template:Cite journal</ref><ref>Template:Cite book</ref>
Home ranges and territoriesEdit
Unlike many other felids, among cheetahs, females tend to occupy larger areas compared to males.<ref name=nowak/> Females typically disperse over large areas in pursuit of prey, but they are less nomadic and roam in a smaller area if prey availability in the area is high. As such, the size of their home range depends on the distribution of prey in a region. In central Namibia, where most prey species are sparsely distributed, home ranges average Template:Cvt, whereas in the woodlands of the Phinda Game Reserve (South Africa), which have plentiful prey, home ranges are Template:Cvt in size.<ref name=hunterwcw/> Cheetahs can travel long stretches overland in search of food; a study in the Kalahari Desert recorded an average displacement of nearly Template:Cvt every day and walking speeds ranged between Template:Cvt.<ref name=marker8/>
Males are generally less nomadic than females; often males in coalitions (and sometimes solitary males staying far from coalitions) establish territories.<ref name=wcw/><ref name=nowak/> Whether males settle in territories or disperse over large areas forming home ranges depends primarily on the movements of females. Territoriality is preferred only if females tend to be more sedentary, which is more feasible in areas with plenty of prey. Some males, called floaters, switch between territoriality and nomadism depending on the availability of females.<ref name=hunterwcw/> A 1987 study showed territoriality depended on the size and age of males and the membership of the coalition.<ref name="caro1987"/> The ranges of floaters averaged Template:Cvt in the Serengeti to Template:Cvt in central Namibia. In the Kruger National Park (South Africa) territories were much smaller. A coalition of three males occupied a territory measuring Template:Cvt, and the territory of a solitary male measured Template:Cvt.<ref name=hunterwcw/> When a female enters a territory, the males will surround her; if she tries to escape, the males will bite or snap at her. Generally, the female can not escape on her own; the males themselves leave after they lose interest in her. They may smell the spot she was sitting or lying on to determine if she was in oestrus.<ref name=marker9/>
CommunicationEdit
The cheetah is a vocal felid with a broad repertoire of calls and sounds; the acoustic features and the use of many of these have been studied in detail.<ref name="volodina">Template:Cite journal</ref> The vocal characteristics, such as the way they are produced, are often different from those of other cats.<ref name="sh2003">Template:Cite journal</ref> For instance, a study showed that exhalation is louder than inhalation in cheetahs, while no such distinction was observed in the domestic cat.<ref>Template:Cite book</ref><ref name=Eklund_al2012>Template:Cite book</ref> Listed below are some commonly recorded vocalisations observed in cheetahs:
- Chirping: A chirp (or a "stutter-bark") is an intense bird-like call and lasts less than a second. Cheetahs chirp when they are excited, for instance, when gathered around a kill. Other uses include summoning concealed or lost cubs by the mother, or as a greeting or courtship between adults.<ref name=sh2003/> The cheetah's chirp is similar to the soft roar of the lion, and its churr as the latter's loud roar.<ref name=volodina/> A similar but louder call ('yelp') can be heard from up to Template:Convert away; this call is typically used by mothers to locate lost cubs, or by cubs to find their mothers and siblings.<ref name=wcw/><ref name="kingdon">Template:Cite book</ref>
- Churring (or churtling): A churr is a shrill, staccato call that can last up to two seconds. Churring and chirping have been noted for their similarity to the soft and loud roars of the lion. It is produced in similar context as chirping, but a study of feeding cheetahs found chirping to be much more common.<ref name=Estes/><ref name=sh2003/>
- Purring: Similar to purring in domestic cats but much louder, it is produced when the cheetah is content, and as a form of greeting or when licking one another.<ref name=wcw/><ref name=sh2003/> It involves continuous sound production alternating between egressive and ingressive airstreams.<ref>Template:Cite book</ref>
- Agonistic sounds: These include bleating, coughing, growling, hissing, meowing and moaning (or yowling). A bleat indicates distress, for instance when a cheetah confronts a predator that has stolen its kill. Growls, hisses and moans are accompanied by multiple, strong hits on the ground with the front paw, during which the cheetah may retreat by a few metres.<ref name=volodina/><ref name=sh2003/><ref name=bare_url>Template:Cite book</ref> A meow, though a versatile call, is typically associated with discomfort or irritation.<ref name=volodina/><ref>Template:Cite journal</ref>
- Other vocalisations: Individuals can make a gurgling noise as part of a close, amicable interaction.<ref name=wcw/> A "nyam nyam" sound may be produced while eating. Apart from chirping, mothers can use a repeated "ihn ihn" is to gather cubs, and a "prr prr" is to guide them on a journey. A low-pitched alarm call is used to warn the cubs to stand still. Bickering cubs can let out a "whirr"—the pitch rises with the intensity of the quarrel and ends on a harsh note.<ref name=Estes/><ref name=sh2003/>
Another major means of communication is by scent—the male will often raise his tail and spray urine on elevated landmarks such as a tree trunks, stumps or rocks; other cheetahs will sniff these landmarks and repeat the ritual. Females may also show marking behaviour but less prominently than males do. Females in oestrus will show maximum urine-marking, and their excrement can attract males from far off. In Botswana, cheetahs are frequently captured by ranchers to protect livestock by setting up traps in traditional marking spots; the calls of the trapped cheetah can attract more cheetahs to the place.<ref name=Estes/><ref name=nowak/>
Touch and visual cues are other ways of signalling in cheetahs. Social meetings involve mutual sniffing of the mouth, anus and genitals. Individuals will groom one another, lick each other's faces and rub cheeks. However, they seldom lean on or rub their flanks against each other. The tear streaks on the face can sharply define expressions at close range. Mothers probably use the alternate light and dark rings on the tail to signal their cubs to follow them.<ref name=Estes/>
Diet and huntingEdit
The cheetah is a carnivore that hunts small to medium-sized prey weighing Template:Cvt, but mostly less than Template:Cvt. Its primary prey are medium-sized ungulates. They are the major component of the diet in certain areas, such as Dama and Dorcas gazelles in the Sahara, impala in the eastern and southern African woodlands, springbok in the arid savannas to the south and Thomson's gazelle in the Serengeti. Smaller antelopes like the common duiker are frequent prey in the southern Kalahari. Larger ungulates are typically avoided, though nyala, whose males weigh around Template:Cvt, were found to be the major prey in a study in the Phinda Game Reserve. In Namibia cheetahs are the major predators of livestock.<ref name=skinner/><ref name=wcw/><ref name="Hayward06">Template:Cite journal</ref> The diet of the Asiatic cheetah consists of chinkara, desert hare, goitered gazelle, urial, wild goats, and livestock; in India cheetahs used to prey mostly on blackbuck.<ref name=hunterwcw/><ref>Template:Cite journal</ref>
Prey preferences and hunting success vary with the age, sex and number of cheetahs involved in the hunt and on the vigilance of the prey. Generally, only groups of cheetahs (coalitions or mother and cubs) will try to kill larger prey; mothers with cubs especially look out for larger prey and tend to be more successful than females without cubs. Individuals on the periphery of the prey herd are common targets; vigilant prey which would react quickly on seeing the cheetah are not preferred.<ref name=Geptner1972>Template:Cite book</ref><ref name=wcw/><ref name=Qumsiyeh1996>Template:Cite book</ref>
Cheetahs are one of the most iconic pursuit predators, hunting primarily throughout the day, sometimes with peaks at dawn and dusk; they tend to avoid larger predators like the primarily nocturnal lion.<ref name=hunterwcw/> Cheetahs in the Sahara and Maasai Mara in Kenya hunt after sunset to escape the high temperatures of the day.<ref name="hunt2">Template:Cite journal</ref> Cheetahs use their vision to hunt instead of their sense of smell; they keep a lookout for prey from resting sites or low branches. The cheetah will stalk its prey, trying to conceal itself in cover, and approach as close as possible, often within Template:Cvt of the prey (or even closer for less alert prey). Alternatively the cheetah can lie hidden in cover and wait for the prey to come nearer. A stalking cheetah assumes a partially crouched posture, with the head lower than the shoulders; it will move slowly and be still at times. In areas of minimal cover, the cheetah will approach within Template:Cvt of the prey and start the chase. The chase typically lasts a minute; in a 2013 study, the length of chases averaged Template:Cvt, and the longest run measured Template:Cvt. The cheetah can give up the chase if it is detected by the prey early or if it cannot make a kill quickly. Being lightly built, cheetahs lack the raw strength to tackle down the prey, and instead catch the prey by performing a kind of foot sweep by hitting the prey's leg or rump with the forepaw or using the strong dewclaw to knock the prey off its balance. Such a fall during a high-speed chase may cause the prey to collapse hard enough to break some of its limbs,<ref name=wcw/><ref name=Estes/> and allow the cheetah to then pounce on the fallen and vulnerable prey.
Cheetahs can decelerate dramatically towards the end of the hunt, slowing down from Template:Cvt to Template:Cvt in just three strides, and can easily follow any twists and turns the prey makes as it tries to flee.<ref name=hunterwcw/> To kill medium- to large-sized prey, the cheetah bites the prey's throat to strangle it, maintaining the bite for around five minutes, within which the prey succumbs to asphyxiation and stops struggling. A bite on the nape of the neck or the snout (and sometimes on the skull) suffices to kill smaller prey.<ref name=wcw/><ref name=Estes/> Cheetahs have an average hunting success rate of 25–40%, higher for smaller and more vulnerable prey.<ref name=hunterwcw/><ref name="O'Brien"/>
Once the hunt is over, the prey is taken near a bush or under a tree; the cheetah, highly exhausted after the chase, rests beside the kill and pants heavily for five to 55 minutes. Meanwhile, cheetahs nearby, who did not take part in the hunt, might feed on the kill immediately. Groups of cheetah consume the kill peacefully, though minor noises and snapping may be observed.<ref name=wcw/> Cheetahs can consume large quantities of food; a cheetah at the Etosha National Park (Namibia) was found to consume as much as Template:Cvt within two hours.<ref>Template:Cite journal</ref> However, on a daily basis, a cheetah feeds on around Template:Cvt of meat.<ref name=Estes/> Cheetahs, especially mothers with cubs, remain cautious even as they eat, pausing to look around for vultures and predators who may steal the kill.<ref>Template:Cite journal</ref>
Cheetahs move their heads from side to side so the sharp carnassial teeth tear the flesh, which can then be swallowed without chewing. They typically begin with the hindquarters where the tissue is the softest, and then progress toward the abdomen and the spine. Ribs are chewed on at the ends, and the limbs are not generally torn apart while eating. Unless the prey is very small, the skeleton is left almost intact after feeding on the meat. Cheetahs might lose up 13–14% of their kills to larger and stronger carnivores.<ref name="Schaller-1972">Template:Cite book</ref> To defend itself or its prey, a cheetah will hold its body low to the ground and snarl with its mouth wide open, the eyes staring threateningly ahead and the ears folded backward. This may be accompanied by moans, hisses and growls, and hitting the ground with the forepaws.<ref name=Estes/> Although uncommon, cases of wild cheetahs scavenging carcasses that not hunt themselves have been observed; even one case of a cheetah stealing a spotted hyena kill is known. Causes of this scavenging behavior are unclear.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>
Cheetahs appear to have a comparatively higher hunting success rate than other predators.<ref name=wcw/> Their success rate for hunting Thomson gazelles is 70%, whereas the success rate of African wild dogs is 57%, of spotted hyenas 33%, and of lions 26%.<ref name=Schaller-1972/> Their success rate for hunting impalas is 26%, but of African wild dogs only 15.5%.<ref name=":1">Template:Cite journal</ref>
Reproduction and life cycleEdit
The cheetah breeds throughout the year; females are polyestrous and induced ovulators with an estrous cycle of 12 days on average that can vary from three days to a month. They have their first litter at two to three years of age and can conceive again after 17 to 20 months from giving birth, or even sooner if a whole litter is lost. Males can breed at less than two years of age in captivity, but this may be delayed in the wild until the male acquires a territory.<ref name=marker9/> A 2007 study showed that females who gave birth to more litters early in their life often died younger, indicating a trade-off between longevity and yearly reproductive success.<ref>Template:Cite journal</ref>
Urine-marking in males can become more pronounced when a female in their vicinity comes into estrus. Males, sometimes even those in coalitions, fight among one another to secure access to the female.<ref>Template:Cite journal</ref> Often one male will eventually win dominance over the others and mate with the female, though a female can mate with different males.<ref>Template:Cite journal</ref> Mating begins with the male approaching the female, who lies down on the ground; individuals often chirp, purr or yelp at this time. No courtship behaviour is observed; the male immediately secures hold of the female's nape, and copulation takes place. The pair then ignore each other, but meet and copulate a few more times three to five times a day for the next two to three days before finally parting ways.<ref name=mammal/><ref name=Estes/><ref>Template:Cite journal</ref>
After a gestation of nearly three months, a litter of one to eight cubs is born (though those of three to four cubs are more common). Births take place at 20–25 minute intervals in a sheltered place such as thick vegetation. The eyes are shut at birth, and open in four to 11 days. Newborn cubs might spit a lot and make soft churring noises; they start walking by two weeks. Their nape, shoulders and back are thickly covered with long bluish-grey hair, called a mantle, which gives them a mohawk-type appearance; this fur is shed as the cheetah grows older.<ref name=Estes/><ref name="caro1994">Template:Cite book</ref> A study suggested that this mane gives a cheetah cub the appearance of a honey badger, and could act as camouflage from attacks by these badgers or predators that tend to avoid them.<ref>Template:Cite journal</ref>
Compared to other felids, cheetah cubs are highly vulnerable to several predators during the first few weeks of their life.<ref name="laurenson2">Template:Cite journal</ref><ref>Template:Cite journal</ref> Mothers keep their cubs hidden in dense vegetation for the first two months and nurse in the early morning. The mother is extremely vigilant at this stage; she stays within Template:Convert of the lair, frequently visits her cubs, moves them every five to six days, and remains with them after dark. Despite trying to make minimal noise, she cannot generally defend her litter from predators. Predation is the leading cause of mortality in cheetah cubs; a study showed that in areas with a low density of predators (such as Namibian farmlands) around 70% of the cubs make it beyond the age of 14 months, whereas in areas like the Serengeti National Park, where several large carnivores exist, the survival rate was just 17%. Deaths also occur from starvation if their mothers abandon them, fires, or pneumonia because of exposure to bad weather.<ref name=Estes/><ref name=marker9/> Generation length of the cheetah is six years.<ref>Template:Cite journal</ref>
Cubs start coming out of the lair at two months of age, trailing after their mother wherever she goes. At this point the mother nurses less and brings solid food to the cubs; they retreat away from the carcass in fear initially, but gradually start eating it. The cubs might purr as the mother licks them clean after the meal. Weaning occurs at four to six months. To train her cubs in hunting, the mother will catch and let go of live prey in front of her cubs.<ref name=marker9/> Cubs' play behaviour includes chasing, crouching, pouncing and wrestling; there is plenty of agility, and attacks are seldom lethal.<ref name=Estes/><ref name=marker9/> Playing can improve catching skills in cubs, though the ability to crouch and hide may not develop remarkably.<ref>Template:Cite journal</ref>
Cubs as young as six months try to capture small prey like hares and young gazelles. However, they may have to wait until as long as 15 months of age to make a successful kill on their own. At around 20 months, offspring become independent; mothers might have conceived again by then. Siblings may remain together for a few more months before parting ways. While females stay close to their mothers, males move farther off.<ref name=Estes/><ref name=marker9/><ref name="Kelly, M. J. 1998">Template:Cite journal</ref> The lifespan of wild cheetahs is 14 to 15 years for females, and their reproductive cycle typically ends by 12 years of age; males generally live as long as ten years.<ref name=iucn />
CompetitionEdit
Template:Expand section Although cheetahs and spotted hyenas favor different prey, the latter will nevertheless steal cheetah kills with no difficulty, with George Schaller observing that cheetahs in the Serengeti lost 4% of their kills to them.<ref name=kruuk>Template:Cite book</ref> Cheetahs, particularly females with cubs, may attempt to protect their kills from hyenas by making threatening vocalizations and lunges, but will retreat if the larger carnivores persist.<ref name=Estes/>
In Iran, cheetahs compete with leopards for chinkara, bezoar ibex and urial. One study undertaken in the Bafq Protected Area found that cheetahs avoided leopards by occupying lower elevations, though one cheetah was nevertheless killed by a leopard during the study.<ref>Template:Cite journal</ref> Cheetah mothers have however been observed to drive off leopards threatening their cubs.<ref name=Estes/>
Distribution and habitatEdit
In eastern and southern Africa, the cheetah occurs mostly in savannas like the Kalahari and Serengeti. In central, northern and western Africa, it inhabits arid mountain ranges and valleys; in the harsh climate of the Sahara, it prefers high mountains, which receive more rainfall than the surrounding desert. The vegetation and water resources in these mountains support antelopes. In Iran, it occurs in hilly terrain of deserts at elevations up to Template:Cvt, where annual precipitation is generally below Template:Cvt; the primary vegetation in these areas is thinly distributed shrubs, less than Template:Cvt tall.<ref name=iucn /><ref name=hunterwcw/><ref name=marker8/>
The cheetah inhabits a variety of ecosystems and appears to be less selective in habitat choice than other felids; it prefers areas with greater availability of prey, good visibility and minimal chances of encountering larger predators. It seldom occurs in tropical forests. It has been reported at the elevation of Template:Cvt. An open area with some cover, such as diffused bushes, is probably ideal for the cheetah because it needs to stalk and pursue its prey over a distance. This also minimises the risk of encountering larger carnivores. The cheetah tends to occur in low densities typically between 0.3 and 3.0 adults per Template:Cvt; these values are 10–30% of those reported for leopards and lions.<ref name=iucn /><ref name=marker8>Template:Cite book</ref>
Historical rangeEdit
In prehistoric times, the cheetah was distributed throughout Africa, Asia and Europe.<ref name=wcw/> It gradually fell to extinction in Europe, possibly because of competition with the lion.<ref name=caro1994/> Today the cheetah has been extirpated in most of its historical range; the numbers of the Asiatic cheetah had begun plummeting since the late 1800s, long before the other subspecies started their decline. As of 2017, cheetahs occur in just nine per cent of their erstwhile range in Africa, mostly in unprotected areas.<ref name=marker4/>
In the past until the mid-20th century, the cheetah ranged across vast stretches in Asia, from the Arabian Peninsula in the west to the Indian subcontinent in the east, and as far north as the Aral and Caspian Seas.<ref name=mallon07>Template:Cite journal</ref> A few centuries ago the cheetah was abundant in India, and its range coincided with the distribution of major prey like the blackbuck.<ref name=wcw/> However, its numbers in India plummeted from the 19th century onward; Divyabhanusinh of the Bombay Natural History Society notes that the last three individuals in the wild were killed by Maharaja Ramanuj Pratap Singh of Surguja in 1947.<ref>Template:Cite journal</ref><ref>Template:Cite news</ref> The last confirmed sighting in India was of a cheetah that drowned in a well near Hyderabad in 1957.<ref name="sharma13">Template:Cite book</ref> In Iran there were around 400 cheetahs before World War II, distributed across deserts and steppes to the east and the borderlands with Iraq to the west; the numbers were falling because of a decline in prey. In Iraq, cheetahs were reported from Basra in the 1920s. Conservation efforts in the 1950s stabilised the population, but prey species declined again in the wake of the Iranian Revolution (1979) and the Iran–Iraq War (1980–1988), leading to a significant contraction of the cheetah's historical range in the region.<ref name=marker4/><ref name=marker5>Template:Cite book</ref>
In 1975, the cheetah population was estimated at 15,000 individuals throughout Sub-Saharan Africa, following the first survey in this region by Norman Myers. The range covered most of eastern and southern Africa, except for the desert region on the western coast of modern-day Angola and Namibia.<ref name=myers>Template:Cite report</ref> In the following years, cheetah populations across the region have become smaller and more fragmented as their natural habitat has been modified dramatically.<ref name=south1>Template:Cite report</ref>
Present distributionEdit
The cheetah occurs mostly in eastern and southern Africa; its presence in Asia is limited to the central deserts of Iran, though there have been unconfirmed reports of sightings in Afghanistan, Iraq and Pakistan in the last few decades.<ref name=iucn /><ref name=marker4/> The global population of cheetahs was estimated at nearly 7,100 mature individuals in 2016. The Iranian population appears to have decreased from 60–100 individuals in 2007 to 43 in 2016, distributed in three subpopulations over less than Template:Cvt in Iran's central plateau.<ref name=durant2017>Template:Cite journal</ref><ref name="hunter2007">Template:Cite journal</ref> The largest population of nearly 4,000 individuals is sparsely distributed over Angola, Botswana, Mozambique, Namibia, South Africa and Zambia. Another population in Kenya and Tanzania comprises about 1,000 individuals. All other cheetahs occur in small, fragmented groups of less than 100 individuals each. Populations are thought to be declining.<ref name=durant2017/>
ThreatsEdit
The cheetah is threatened by several factors, like habitat loss and fragmentation of populations. Habitat loss is caused mainly by the introduction of commercial land use, such as agriculture and industry.<ref name=iucn /> It is further aggravated by ecological degradation, like woody plant encroachment, which is common in southern Africa.<ref>Template:Cite journal</ref><ref name=FSC/><ref>Template:Cite journal</ref> Moreover, the species apparently requires a sizeable area to live in as indicated by its low population densities. Shortage of prey and conflict with other species such as humans and large carnivores are other major threats.<ref name=iucn /><ref name=marker10>Template:Cite book</ref> The cheetah appears to be less capable of coexisting with humans than the leopard.<ref>Template:Cite journal</ref> With 76% of its range consisting of unprotected land, the cheetah is often targeted by farmers and pastoralists who attempt to protect their livestock, especially in Namibia.<ref>Template:Cite journal</ref> Illegal wildlife trade and trafficking is another problem in some places (like Ethiopia). Some tribes, like the Maasai people in Tanzania, have been reported to use cheetah skins in ceremonies.<ref name=marker1/><ref name=marker2/> Roadkill is another threat, especially in areas where roads have been constructed near natural habitat or protected areas. Cases of roadkill involving cheetahs have been reported from Kalmand, Touran National Park, and Bafq in Iran.<ref name=iucn /> The reduced genetic variability makes cheetahs more vulnerable to diseases;<ref name=obrien2017/> however, the threat posed by infectious diseases may be minor, given the low population densities and hence a reduced chance of infection.<ref name=iucn />
ConservationEdit
The cheetah has been classified as Vulnerable on the IUCNRed List; it is listed under AppendixTemplate:NbspI of the Convention on the Conservation of Migratory Species of Wild Animals and AppendixTemplate:NbspI of the Convention on International Trade in Endangered Species.<ref name=iucn /> The Endangered Species Act enlists the cheetah as Endangered.<ref name=ecos>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
In AfricaEdit
Until the 1970s, cheetahs and other carnivores were frequently killed to protect livestock in Africa. Gradually the understanding of cheetah ecology increased and their falling numbers became a matter of concern. The De Wildt Cheetah and Wildlife Centre was set up in 1971 in South Africa to provide care for wild cheetahs regularly trapped or injured by Namibian farmers.<ref name=marker1/> By 1987, the first major research project to outline cheetah conservation strategies was underway.<ref>Template:Cite journal</ref> The Cheetah Conservation Fund, founded in 1990 in Namibia, put efforts into field research and education about cheetahs on the global platform.<ref name=marker1/> The CCF runs a cheetah genetics laboratory, the only one of its kind, in Otjiwarongo (Namibia);<ref>Template:Cite news</ref> "Bushblok" is an initiative to restore habitat systematically through targeted bush thinning and biomass utilisation.<ref name=FSC>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> Several more cheetah-specific conservation programmes have since been established, like Cheetah Outreach in South Africa.<ref name=marker1/>
The Global Cheetah Action Plan Workshop in 2002 laid emphasis on the need for a range-wide survey of wild cheetahs to demarcate areas for conservation efforts and on creating awareness through training programs.<ref>Template:Cite report</ref> The Range Wide Conservation Program for Cheetah and African Wild Dogs began in 2007 as a joint initiative of the IUCN Cat and Canid Specialist Groups, the Wildlife Conservation Society and the Zoological Society of London. National conservation plans have been developed for several African countries.<ref>Template:Cite news</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2014, the CITES Standing Committee recognised the cheetah as a "species of priority" in their strategies in northeastern Africa to counter wildlife trafficking.<ref name=cites>Template:Cite report</ref> In December 2016, the results of an extensive survey detailing the distribution and demography of cheetahs throughout the range were published; the researchers recommended listing the cheetah as Endangered on the IUCN Red List.<ref name=durant2017/>
The cheetah was reintroduced in Malawi in 2017.<ref>Template:Cite news</ref>
In AsiaEdit
In 2001, the Iranian government collaborated with the CCF, the IUCN, Panthera Corporation, United Nations Development Programme and the Wildlife Conservation Society on the Conservation of Asiatic Cheetah Project (CACP) to protect the natural habitat of the Asiatic cheetah and its prey.<ref name="CatWatch">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2004, the Iranian Centre for Sustainable Development (CENESTA) conducted an international workshop to discuss conservation plans with local stakeholders.<ref name=marker1/> Iran declared 31Template:NbspAugust as National Cheetah Day in 2006.<ref>Template:Cite news</ref> The Iranian Cheetah Strategic Planning meet in 2010 formulated a five-year conservation plan for Asiatic cheetahs.<ref name=marker1/> The CACP Phase II was implemented in 2009, and the third phase was drafted in 2018.<ref>Template:Cite news</ref>
During the early 2000s scientists from the Centre for Cellular and Molecular Biology (Hyderabad) proposed a plan to clone Asiatic cheetahs from Iran for reintroduction in India, but Iran denied the proposal.<ref>Template:Cite news</ref> In September 2009, the Minister of Environment and Forests assigned the Wildlife Trust of India and the Wildlife Institute of India with examining the potential of importing African cheetahs to India.<ref>Template:Cite news</ref> Kuno Wildlife Sanctuary and Nauradehi Wildlife Sanctuary were suggested as reintroduction sites for the cheetah because of the high prey density.<ref>Template:Cite report</ref> However, plans for reintroduction were stalled in May 2012 by the Supreme Court of India because of a political dispute and concerns over introducing a non-native species to the country. Opponents stated the plan was "not a case of intentional movement of an organism into a part of its native range".<ref>Template:Cite news</ref><ref>Template:Cite magazine</ref> On 28 January 2020, the Supreme Court allowed the central government to introduce cheetahs to a suitable habitat in India on an experimental basis to see if they can adapt to it.<ref>Template:Cite newsTemplate:Cbignore</ref><ref>Template:Cite magazine</ref> In 2020, India signed a memorandum of understanding with Namibia as part of Project Cheetah.<ref>Template:Cite news</ref> In July 2022, it was announced that eight cheetahs would be transferred from Namibia to India in August.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The eight cheetahs were released into Kuno National Park on 17 September 2022.<ref>Template:Cite news</ref> Since their introduction, they gave birth to 17 cubs. However, by September 2024, eight adult cheetahs and four cubs already died.<ref>Template:Cite news</ref>
Interaction with humansEdit
TamingEdit
The cheetah shows little aggression toward humans, and can be tamed easily, as it has been since antiquity.<ref name=caro1994/> The earliest known depictions of the cheetah are from the Chauvet Cave in France, dating back to 32,000–26,000 BC.<ref name="marker2alt">Template:Cite book</ref> According to historians such as Heinz Friederichs and Burchard Brentjes, the cheetah was first tamed in Sumer and this gradually spread out to central and northern Africa, from where it reached India. The evidence for this is mainly pictorial; for instance, a Sumerian seal dating back to Template:Circa, featuring a long-legged leashed animal has fueled speculation that the cheetah was first tamed in Sumer. However, Thomas Allsen argues that the depicted animal might be a large dog.<ref name="allsen">Template:Cite book</ref> Other historians, such as Frederick Zeuner, have opined that ancient Egyptians were the first to tame the cheetah, from where it gradually spread into central Asia, Iran and India.<ref name="mair">Template:Cite book</ref>
In comparison, theories of the cheetah's taming in Egypt are stronger and include timelines proposed on this basis.<ref name=mair/> Mafdet, one of the ancient Egyptian deities worshiped during the First Dynasty (3100–2900Template:NbspBC), was sometimes depicted as a cheetah. Ancient Egyptians believed the spirits of deceased pharaohs were taken away by cheetahs.<ref name=marker2alt/> Reliefs in the Deir el-Bahari temple complex tell of an expedition by Egyptians to the Land of Punt during the reign of Hatshepsut (1507–1458Template:NbspBC) that fetched, among other things, animals called "panthers". During the New Kingdom (16th to 11th centuries BC), cheetahs were common pets for royalty, who adorned them with ornate collars and leashes.<ref name=mair/> Rock carvings depicting cheetahs dating back to 2000–6000 years ago have been found in Twyfelfontein; little else has been discovered in connection to the taming of cheetahs (or other cats) in southern Africa.<ref name=marker2alt/>
Hunting cheetahs are known in pre-Islamic Arabic art from Yemen.<ref>Template:Cite journal</ref> Hunting with cheetahs became more prevalent toward the seventh centuryTemplate:NbspAD. In the Middle East, the cheetah would accompany the nobility to hunts in a special seat on the back of the saddle. Taming was an elaborate process and could take a year to complete.<ref name=marker2alt/> The Romans may have referred to the cheetah as the Template:Transliteration ({{#invoke:Lang|lang}}) or Template:Transliteration ({{#invoke:Lang|lang}}), believing it to be a hybrid between a leopard and a lion because of the mantle seen in cheetah cubs and the difficulty of breeding them in captivity.<ref name="Nicholas-1999">Template:Cite journal</ref> A Roman hunting cheetah is depicted in a 4th-century mosaic from Lod, Israel.<ref>Template:Cite journal</ref> Cheetahs continued to be used into the Byzantine period of the Roman Empire, with "hunting leopards" being mentioned in the Cynegetica (283/284 AD).<ref name="Nicholas-1999" /><ref>Template:Cite book</ref><ref>Template:Cite book</ref>
In eastern Asia, records are confusing as regional names for the leopard and the cheetah may be used interchangeably. The earliest depiction of cheetahs from eastern Asia dates back to the Tang dynasty (7th to 10th centuriesTemplate:NbspAD); paintings depict tethered cheetahs and cheetahs mounted on horses. Chinese emperors would use cheetahs and caracals as gifts. In the 13th and 14th centuries, the Yuan rulers bought numerous cheetahs from the western parts of the empire and from Muslim merchants. According to the Template:Transliteration, the subsequent Ming dynasty (14th to 17th centuries) continued this practice.<ref name=mair/> Tomb figurines from the Mongol empire, dating back to the reign of Kublai Khan (1260–1294Template:NbspAD), represent cheetahs on horseback.<ref name=marker2alt/> The Mughal ruler Akbar the Great (1556–1605Template:NbspAD) is said to have kept as many as 1000 khasa (imperial) cheetahs.<ref name="O'Brien" /><ref name=marker2alt/> His son Jahangir wrote in his memoirs, Tuzk-e-Jahangiri, that only one of them gave birth.<ref name=mair/> Mughal rulers trained cheetahs and caracals in a similar way as the western Asians, and used them to hunt game, especially blackbuck. The rampant hunting severely affected the populations of wild animals in India; by 1927, cheetahs had to be imported from Africa.<ref name=marker2alt/>
In captivityEdit
The first cheetah to be brought into captivity in a zoo was at the Zoological Society of London in 1829. Early captive cheetahs showed a high mortality rate, with an average lifespan of 3–4 years. After trade of wild cheetahs was delimited by the enforcement of CITES in 1975, more efforts were put into breeding in captivity; in 2014 the number of captive cheetahs worldwide was estimated at 1730 individuals, with 87% born in captivity.<ref name=marker1/><ref name=marker22>Template:Cite book</ref>
Mortality under captivity is generally high; in 2014, 23% of the captive cheetahs worldwide died under one year of age, mostly within a month of birth.<ref name=marker22/> Deaths result from several reasons—stillbirths, birth defects, cannibalism, hypothermia, maternal neglect, and infectious diseases.<ref>Template:Cite journal</ref> Compared to other felids, cheetahs need specialised care because of their higher vulnerability to stress-induced diseases; this has been attributed to their low genetic variability and factors of captive life.<ref name=marker25>Template:Cite book</ref> Common diseases of cheetahs include feline herpesvirus, feline infectious peritonitis, gastroenteritis, glomerulosclerosis, leukoencephalopathy, myelopathy, nephrosclerosis and veno-occlusive disease.<ref name=marker25/><ref name=munson>Template:Cite journal</ref> High density of cheetahs in a place, closeness to other large carnivores in enclosures, improper handling, exposure to public and frequent movement between zoos can be sources of stress for cheetahs. Recommended management practices for cheetahs include spacious and ample access to outdoors, stress minimisation by exercise and limited handling, and following proper hand-rearing protocols (especially for pregnant females).<ref name=marker24>Template:Cite book</ref>
Wild cheetahs are far more successful breeders than captive cheetahs;<ref>Template:Cite journal</ref> this has also been linked to increased stress levels in captive individuals.<ref name=marker25/> In a study in the Serengeti, females were found to have a 95% success rate in breeding, compared to 20% recorded for North American captive cheetahs in another study.<ref name=fcr>Template:Cite journal</ref><ref>Template:Cite journal</ref> On 26 November 2017, a female cheetah gave birth to eight cubs at the Saint Louis Zoo, setting a record for the most births recorded by the Association of Zoos and Aquariums.<ref>Template:Cite news</ref> Chances of successful mating in captive males can be improved by replicating social groups such as coalitions observed in the wild.<ref>Template:Cite journal</ref>
Attacks on humansEdit
There are no documented records of lethal attacks on humans by wild cheetahs.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> However, there have been instances of people being fatally mauled by captive cheetahs. In 2007, a 37-year-old woman from Antwerp was killed by a cheetah in a Belgian zoo after sneaking into its cage outside of visiting hours.<ref>Woman killed by cheetah in Belgian zoo Template:Webarchive. Sydney Morning Herald. 13 February 2007. Retrieved 23 October 2023</ref> In 2017, a three-year-old child was attacked by a captive cheetah on a farm in Philippolis, South Africa. Despite being airlifted to a hospital in Bloemfontein, the boy died from his injuries.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
In cultureEdit
The cheetah has been widely portrayed in a variety of artistic works. In Bacchus and Ariadne, an oil painting by the 16th-century Italian painter Titian, the chariot of the Greek god Dionysus (Bacchus) is depicted as being drawn by two cheetahs. The cheetahs in the painting were previously considered to be leopards.<ref>Template:Cite journal</ref> In 1764, English painter George Stubbs commemorated the gifting of a cheetah to George III by the English Governor of Madras, Sir George Pigot in his painting Cheetah with Two Indian Attendants and a Stag. The painting depicts a cheetah, hooded and collared by two Indian servants, along with a stag it was supposed to prey upon.<ref>Template:Cite book</ref> The 1896 painting The Caress by the 19th-century Belgian symbolist painter Fernand Khnopff is a representation of the myth of Oedipus and the Sphinx and portrays a creature with a woman's head and a cheetah's body.<ref>Template:Cite book</ref>
Two cheetahs are depicted standing upright and supporting a crown in the coat of arms of the Free State (South Africa).<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
In 1969, Joy Adamson, of Born Free fame, wrote The Spotted Sphinx, a biography of her pet cheetah Pippa.<ref>Template:Cite book</ref> Hussein, An Entertainment, a novel by Patrick O'Brian set in the British Raj period in India, illustrates the practice of royalty keeping and training cheetahs to hunt antelopes.<ref>Template:Cite book</ref> The book How It Was with Dooms tells the true story of a family raising an orphaned cheetah cub named Dooms in Kenya. The 2005 film Duma was based loosely on this book.<ref>Template:Cite book</ref> The animated series ThunderCats had a character named "Cheetara", an anthropomorphic cheetah, voiced by Lynne Lipton.<ref>Template:Cite book</ref> Comic book heroine Wonder Woman's chief adversary is Barbara Ann Minerva alias The Cheetah.<ref>Template:Cite book</ref>
The Bill Thomas Cheetah American racing car, a Chevrolet-based coupe first designed and driven in 1963, was never homologated for competition beyond prototype status; its production ended in 1966.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 1986, Frito-Lay introduced Chester Cheetah, an anthropomorphic cheetah, as the mascot for their snack food Cheetos.<ref>Template:Cite book</ref> The Mac OS X 10.0 was code-named "Cheetah".<ref>Template:Cite news</ref>
See alsoEdit
ReferencesEdit
Further readingEdit
External linksEdit
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