Felidae

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Felidae[1]
Temporal range:
OligocenePresent, 25–0 Ma
The Felidae.jpg
Clockwise from top left: tiger (Panthera tigris), Canadian lynx (Lynx canadensis), fishing cat (Prionailurus viverrinus), wildcat (Felis silvestris), ocelot (Leopardus pardalis), Asian golden cat (Catopuma temminckii), serval (Leptailurus serval) and cougar (Puma concolor).
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Fischer von Waldheim, 1817
Type genus
Felis
Subfamilies

Pantherinae
Felinae
Machairodontinae
Proailurinae[2]

Felidae range.png
Felidae ranges

Felidae is a family of mammals in the order Carnivora, colloquially referred to as cats. A member of this family is also called a felid.[3][4][5][6] The term "cat" refers both to felids in general and specifically to the domestic cat (Felis catus).[7]

Reginald Innes Pocock divided the extant Felidae into three subfamilies: the Pantherinae, the Felinae and the Acinonychinae, differing from each other by the ossification of the hyoid apparatus and by the cutaneous sheaths which protect their claws.[8] This concept has been revised following developments in molecular biology and techniques for analysis of morphological data. Today, the living Felidae are divided in two subfamilies, with the Pantherinae including seven Panthera and two Neofelis species. The Felinae include all the non-pantherine cats with 10 genera and 34 species.[9]

The first cats emerged during the Oligocene, about 25 million years ago, with the appearance of Proailurus and Pseudaelurus. The latter species complex was ancestral to two main lines of felids: the cats in the extant subfamilies and a third major group of extinct cats of the subfamily Machairodontinae. The machairodonts included the saber-toothed cats such as the Smilodon. The "false sabre toothed cats", the Barbourofelidae and Nimravidae, are not true cats, but are closely related and together with Felidae and other cat-like carnivores (hyaenas, viverrids and mongooses) make up the feliform carnivores.The characteristic features of cats have evolved to support a carnivorous lifestyle, with adaptations for ambush or stalking and short pursuit hunting.[7] They have slender muscular bodies, strong flexible forelimbs and retractable claws for holding prey, dental and cranial adaptations for a strong bite, and often have characteristic striped or spotted coat patterns for camouflage.

Characteristics[edit]

All members of the cat family have the following characteristics in common:

  • They are digitigrade, have five toes on their forefeet and four on their hind feet. Their curved claws are protractile and attached to the terminal bones of the toe with ligaments and tendons. The claws are guarded by cutaneous sheaths, except in the Acinonyx.[10]
  • They actively protract the claws by contracting muscles in the toe,[11] and they passively retract them. The dewclaws are expanded but do not protract.[12]
  • They have 30 teeth with a dental formula of 3.1.3.13.1.2.1. The upper third premolar and lower molar are adapted as carnassial teeth, suited to tearing and cutting flesh.[13] The canine teeth are large, reaching exceptional size in the extinct saber-toothed species. The lower carnassial is smaller than the upper carnassial and has a crown with two compressed blade-like pointed cusps.[11]
  • Their nose projects slightly beyond the lower jaw.[10]
  • They have well developed and highly sensitive whiskers above the eyes, on the cheeks, on the muzzle, but not below the chin.[10] Whiskers help to navigate in the dark and to capture and hold prey.[12]
  • Their skull is foreshortened with a rounded profile and large orbits.[12]
  • Their tongue is covered with horny papillae, which rasp meat from prey and aid in grooming.[12]
  • Their eyes are relatively large, situated to provide binocular vision. Their night vision is especially good due to the presence of a tapetum lucidum, which reflects light back inside the eyeball, and gives felid eyes their distinctive shine. As a result, the eyes of felids are about six times more light sensitive than those of humans, and many species are at least partially nocturnal. The retina of felids also contains a relatively high proportion of rod cells, adapted for distinguishing moving objects in conditions of dim light, which are complemented by the presence of cone cells for sensing colour during the day.[11]
  • Their external ears are large, and especially sensitive to high-frequency sounds in the smaller cat species. This sensitivity allows them to locate small rodent prey.[11]
  • They have lithe and flexible bodies with muscular limbs.[11]
  • The plantar pads of both fore and hind feet form compact three-lobed cushions.[13]
  • The penis is subconical and boneless.[10] Relative to body size, they have shorter bacula than canids.[14]

The colour, length and density of their fur are highly variable. Fur colour varies from brown to golden, and fur pattern from distinctive small spots, stripes, to small blotches and rosettes. Those living in cold environments have thick fur with long hair, like the snow leopard and the Pallas's cat.[12] Those living in tropical and hot climate zones have short fur. The only cat species lacking significant markings are the lion, cougar, caracal, jungle cat and jaguarundi. Several species exhibit melanism with all-black individuals.[11]

In the great majority of species, the tail is between a third and a half of the body length, although with some exceptions, like the Lynx species and margay.[11] Cat species vary greatly in body and skull sizes, and weights:

  • The largest cat species is the tiger, with a head-to-body length of up to 390 cm (150 in), a weight range of at least 65 to 325 kg (143 to 717 lb), and a skull length ranging from 316 to 413 mm (12.4 to 16.3 in).[11][15] Although the maximum skull length of a lion is slightly greater at 419 mm (16.5 in), it is generally smaller in head-to-body length than the former.[16]
  • The smallest cat species are the rusty-spotted cat and the black-footed cat. The former is 35 to 48 cm (14 to 19 in) in length and weighs 0.9 to 1.6 kg (2.0 to 3.5 lb).[11] The latter has a head-to-body length of 36.7 to 43.3 cm (14.4 to 17.0 in) and a maximum recorded weight of 2.45 kg (5.4 lb).[17][18]

Senses[edit]

Felids also have a highly developed sense of smell, although not to the degree seen in canids; this is further supplemented by the presence of a vomeronasal organ in the roof of the mouth, allowing the animal to "taste" the air. The use of this organ is associated with the Flehmen response, in which the upper lip is curled upwards. Most felids are unable to taste sweetness due to a mutated gene in their taste buds.

Most felids are able to land on their feet after a fall due to the cat righting reflex.

Vocalisations[edit]

All felids share a broadly similar set of vocalisations, but with some variation between species. In particular, the pitch of calls varies, with larger species producing deeper sounds.

All felids are able to spit, hiss, growl, snarl, and mew. The first four sounds are all used in an aggressive context. The spitting sound is a sudden burst, typically used when making threats, especially towards other species. The hiss is a prolonged, atonal sound used in close range to other members of the species, when the animal is uncertain whether to attack or retreat.

The mewing sound may be used either as a close-contact call, typically between a mother and kittens, or as a louder, longer distance call, primarily during the mating season. The acoustic properties of the mew vary somewhat between different felid species; extreme examples include the whistling sound made by cougars and the mew-grunt of lions and tigers.[citation needed]

Domestic cat purring and meowing.

Most felids seem to be able to purr, vibrating the muscles in their larynx to produce a distinctive buzzing sound. In the wild, purring is used while a mother is caring for kittens. Precisely which species of felids are able to purr is a matter of debate, but the sound has been recorded in most of the smaller species, as well as being common for the cheetah and cougar, and may also be found in other big cats.

Other common felid vocalisations include the gurgle, wah-wah, prusten, and roar. The first two sounds are found only among the Felinae (small cats). Gurgling is a quiet sound used during meetings between friendly individuals, as well as during courtship and when nursing kittens. The wah-wah is a short, deep-sounding call used in close contact, and is not found in all species (it is, for example, absent in the domestic cat).

In contrast, only Panthera species can prusten and roar. Prusten is a short, soft, snorting sound reported in tigers, jaguars, snow leopards, and clouded leopards; it is used during contact between friendly individuals. The roar is an especially loud call with a distinctive pattern that depends on the species. The ability to roar comes from an elongated and specially adapted larynx and hyoid apparatus.[19] When air passes through the larynx on the way from the lungs, the cartilage walls of the larynx vibrate, producing sound. Only lions, leopards, tigers, and jaguars are truly able to roar, although the loudest mews of snow leopards have a similar, if less structured, sound.[11]

Classification[edit]

Traditionally, five subfamilies have been distinguished within the Felidae based on phenotypical features: the Pantherinae, the Felinae, the Acinonychinae[8], and the extinct Machairodontinae and Proailurinae.[2]

Extant species[edit]

Molecular phylogenetic analysis indicates that living felids fall into eight lineages (clades).[20][21] The lineages 5 to 8 are more related to each other than to any of the lineages 1 to 4, so form a clade within the Felinae.[22]

The following is the complete list of genera within the Felidae, grouped according to the traditional phenotypical classification with the corresponding eight genotypical lineages indicated.[22][9]

Subfamily Pantherinae
Genus Species IUCN Red List status and distribution
Panthera [Lineage 1] Tiger (P. tigris) (Linnaeus, 1758)[23]

Panthera tigris tigris.jpg

EN[24]

Tiger map.jpg

Lion (P. leo) (Linnaeus, 1758)[25]

Lion waiting in Namibia.jpg

VU[26]

Lion distribution.png

Jaguar (P. onca) (Linnaeus, 1758)[27]

Standing jaguar.jpg

NT[28]

Panthera onca distribution.svg

Leopard (P. pardus) (Linnaeus, 1758)[29]

Leopard (Panthera pardus).jpg

VU[30]

Leopard distribution.jpg

Snow leopard (P. uncia) (Schreber, 1775)[31]

Schneeleopard Koeln.jpg

VU[32]

Snow leopard range.png

Neofelis [Lineage 1] Clouded leopard (N. nebulosa) (Griffith, 1821)[33]

Neofelis nebulosa.jpg

VU[34]

Clouded-leopard distribution.jpg

Sunda clouded leopard (N. diardi) (Cuvier, 1823)[35]

Borneo clouded leopard.jpg

VU[36]

Sunda-Clouded-leopard distribution.jpg

Subfamily Felinae
Genus Species IUCN Red List status and distribution
Catopuma [Lineage 2] Asian golden cat (Catopuma temminckii) (Vigors & Horsfield, 1827)[37]

Asian golden cat at Edingburgh Zoo.jpg

NT[38]

AsianGoldenCat distribution.jpg

Bay cat (Catopuma badia) (Gray, 1874)[39]

Bay cat 1 Jim Sanderson-cropped.jpg

EN[40]

Bay cat distribution map.png

Pardofelis [Lineage 2] Marbled cat (P. marmorata) (Martin, 1836)[41]

Marbled cat borneo.jpg

NT[42]

MarbledCat distribution.jpg

Caracal [Lineage 3] Caracal (C. caracal) (Schreber, 1776)[43]

Caracl (01), Paris, décembre 2013.jpg

LC[44]

Caracal distribution.jpg

African golden cat (C. aurata) (Temminck, 1827)[45]

FelisAurataKeulemans.jpg

VU[46]

AfricanGoldenCat distribution.jpg

Leptailurus [Lineage 3] Serval (L. serval) (Schreber, 1775)[47]

Leptailurus serval -Serengeti National Park, Tanzania-8.jpg

LC[48]

Serval distribution.jpg

Leopardus [Lineage 4] Ocelot (L. pardalis) (Linnaeus, 1758)[49]

Ocelot (Jaguatirica) Zoo Itatiba.jpg

LC[50]

Ocelot area.png

Oncilla (L. tigrinus) (Schreber, 1775)[51]

Leopardus tigrinus - Parc des Félins.jpg

VU[52]

Oncilla area.png

Pampas cat (L. colocola) (Molina, 1782)[53]

Leopardus pajeros 20101006.jpg

NT[54]

Leopardus colocolo range map.png

Kodkod (L. guigna) (Molina, 1782)[53]

Leopardus guigna.jpeg

VU[55]

Oncifelis guigna dis.png

Margay (L. wiedii) (Schinz, 1821)[56]

Margaykat Leopardus wiedii.jpg

NT[57]

Margay area.png

Geoffroy's cat (L. geoffroyi) (d'Orbigny & Gervais, 1844)[58]

Salzkatze.jpg

LC[59]

Leopardus geoffroyi range map.png

Andean mountain cat (L. jacobitus) (Cornalia, 1865)[60]

Andean cat 1 Jim Sanderson.jpg

EN[61]

AndeanCat distribution.jpg

Southern tigrina (L. guttulus) (Hensel, 1872)[62] VU[63]

Leopardus guttulus range map.png

Lynx [Lineage 5] Eurasian lynx (L. lynx) (Linnaeus, 1758)[64]

Lynx Nationalpark Bayerischer Wald 01.jpg

LC[65]

Eurasian Lynx area.png

Bobcat (L. rufus) Schreber, 1777[66]

Bobcat2.jpg

LC[67]

Bobcat Lynx rufus distribution map.png

Canada lynx (L. canadensis) Kerr, 1792[68]

Lynx-canadensis.jpg

LC[69]

Canada Lynx area.png

Iberian lynx (L. pardinus) (Temminck, 1827)[70]

Linces19.jpg

EN[71]

Mapa distribuicao lynx pardinus defasado.png

Acinonyx [Lineage 6] Cheetah (A. jubatus) Schreber, 1775)[72]

Cheetah (Acinonyx jubatus) female 2.jpg

VU[73]

Cheetah range - 2.png

Puma [Lineage 6] Cougar (P. concolor) Linnaeus, 1771[74]

Mountain Lion in Glacier National Park.jpg

LC[75]

Cougar range map 2010.png

Jaguarundi (Herpailurus yagouaroundi) (Geoffroy Saint-Hilaire, 1803)[76]

Puma yagouaroundi.jpg

LC[77]

Jaguarundi area.png

Otocolobus [Lineage 7] Pallas's cat (O. manul) (Pallas, 1776)[78]

Manoel.jpg

NT[79]

Manul distribution.jpg

Prionailurus [Lineage 7] Leopard cat (P. bengalensis) (Kerr, 1792)[80]

Close-up of a Leopard Cat in Sundarban.jpg

LC[81]

LeopardCat distribution.jpg

Sunda leopard cat (P. javanensis) (Desmarest, 1816)[82]

Blacan Indonesia.jpg

SundaLeopardCat distribution.jpg

Flat-headed cat (P. planiceps) (Vigors & Horsfield, 1827)[37]

Flat-headed cat 1 Jim Sanderson.JPG

VU[83]

Plionailurus planiceps former distribution.png

Fishing cat (P. viverrinus) (Bennett, 1833)

Fishing Cat (120780371).jpeg

VU[84]

FishingCat distribution.jpg

Rusty-spotted cat (P. rubiginosus) (Geoffroy Saint-Hilaire, 1834)[85]

Rusty spotted cat 1.jpg

NT[86]

Prionailurus rubiginosus range map.png

Felis [Lineage 8] Domestic cat (F. catus) Linnaeus, 1758[87]

Jammlich crop.jpg

European wildcat (F. silvestris) Schreber, 1777[88]

European Wildcat Nationalpark Bayerischer Wald 03.jpg

LC[89]

EuropeanWildcat distribution.jpg

Jungle cat (F. chaus) Schreber, 1777[90]

Jungle Cat Felis chaus by Dr. Raju Kasambe DSCN7957 (3).jpg

LC[91]

Distribution of Jungle Cat.jpg

African wildcat (F. lybica) Forster, 1780[92]

Parc des Felins Chat de Gordoni 28082013 2.jpg

AfricanWildcat distribution.jpg

Black-footed cat (F. nigripes) Burchell, 1824[93]

Blackfooted2.jpg

VU[94]

Black-footedCat distribution.jpg

Sand cat (F. margarita) Loche, 1858[95]

Persian sand CAT.jpg

LC[96]

SandCat distribution.jpg

Chinese mountain cat (F. bieti) Milne-Edwards, 1892[97]

Chinese Mountain Cat (Felis Bieti) in XiNing Wild Zoo.jpg

VU[98]

ChineseMountainCat distribution.jpg

Fossil genera[edit]

The American lion was one of the abundant Pleistocene megafauna, a wide variety of very large mammals that became extinct about 10,000 years ago.[99]

The list follows McKenna and Bell's Classification of Mammals for prehistoric genera. The list differs from McKenna and Bell as follows: Sivapanthera is included in the Felinae, as Acinonychinae is no longer recognised as distinct subfamily; Viretailurus is considered a synonym of Puma; Ischyrosmilus is considered a synonym of the genus Homotherium;[100] and several newly recognised genera, including Miracinonyx, Lokotunjailurus and Xenosmilus, have been added.

Fossil felids[edit]

Possibly the oldest known true felid (Proailurus) lived in the late Oligocene and early Miocene epochs. During the Miocene, it gave way to Pseudaelurus. Pseudaelurus is believed to be the latest common ancestor of the two extant subfamilies and the extinct subfamily, Machairodontinae. This group, better known as the saber-tooth cats, became extinct in the Late Pleistocene era. The group includes the genera Smilodon, Machairodus and Homotherium. The Metailurini were originally classified as a distinct tribe within Machairodontinae, though they count as members of the Felinae in recent times.[105][106] Most extinct cat-like animals, once regarded as members of the Felidae, later turned out to be members of related, but distinct, families: the "false sabretooths" Nimravidae and Barbourofelidae. As a result, sabretooth "cats" seem to belong to four different lineages. The total number of fossil felids known to science is low compared to other carnivoran families, such as dogs and bears. Felidae radiated quite recently and most of the extant species are relatively young.

Evolution[edit]

Feliform evolutionary timeline

Results of mitochondrial analysis indicates that all the Felidae descended from a common ancestor. Cats originated in Asia and spread across continents by crossing land bridges. Testing of mitochondrial and nuclear DNA revealed that the ancient cats evolved into eight main lineages that diverged in the course of at least 10 migrations (in both directions) from continent to continent via the Bering land bridge and the Isthmus of Panama, with the genus Panthera being the oldest and the genus Felis being the youngest. About 60% of the modern cat species are estimated to have developed within the last million years.[22]

The Felidae's closest relatives are thought to be the Asiatic linsangs.[107] Together with the viverrids, hyenas, mongooses, and Malagasy carnivores, they form the suborder Feliformia.[108]

Most cat species share a genetic anomaly that prevents them from tasting sweetness.[109] This is believed to be because their diet consists so strictly of meat (which contains no concentrated sugar) and therefore the taste of sugar signals nothing important for the felid brain and confers no evolutionary advantages. While some individual felids may be able to taste sugar, this ability is not characteristic of any felid species as a whole.

Most cat species have a haploid number of 18 or 19. New World cats (those in Central and South America) have a haploid number of 18, possibly due to the combination of two smaller chromosomes into a larger one.[110]

Domestic cats may either have a long or short tail. At one point, biologists had to consider whether the short tail also found in the lynx was the ancestral or derived trait. Without looking at the fossil record, researchers were able to look at the character states found in their outgroups. Because all animals belonging to Felidae's sister taxa, Viverridae, have long tails, scientists could infer that this character state represents the ancestral trait.[108]

Some domestic cats display a rosette pattern on their coats. This character state, however, is not related to the rosettes found on big cats. Domestic cats and big cats underwent convergent evolution for this trait. The most common ancestor to all cats had a flecked coat. Lynxes display this character state. The jaguarundi lost this character state secondarily. The most common recent ancestor of snow leopards, tigers, jaguars, lions, and leopards developed a coat with rosette patterns from the flecked patterns. Tigers and lions, however, do not display rosettes as adults. They both have lost this ancestral character state over time. Adult tigers actually display elongated rosettes that now appear as stripes. Adult lions seem to lack any distinctive markings altogether. Both juvenile tigers and lions, however, display partial rosettes. This ancestral character state appears only during these early stages, supporting the notion that ontogeny reflects phylogeny. The rosette patterns found on snow leopards, jaguars, and leopards all have a common origin.[111]

Fossil occurrences indicate that the Felidae arrived in North America about 10 million years later than the Canidae, and about 20 million years later than the Ursidae and the Nimravidae.[112]

Phylogeny[edit]

The phylogenetic relationships of extant felids are shown in the following cladogram, based on the molecular phylogenetic analysis of Johnson et al. (2006).[22] The lineages, genera and species are as used in that study.

Felidae
Felidae
Panthera lineage
Pantherinae
Neofelis

Neofelis nebulosa (clouded leopard)

Neofelis diardi (Sunda clouded leopard)

Panthera

Panthera uncia (snow leopard)

Panthera tigris (tiger)

Panthera onca (jaguar)

Panthera pardus (leopard)

Panthera leo (lion)

Felinae
Bay cat lineage
Pardofelis

Pardofelis marmorata (marbled cat)

Catopuma

Catopuma badia (bay cat)

Catopuma temminckii (Asian golden cat)

Caracal lineage
Leptailurus

Leptailurus serval (serval)

Caracal

Caracal caracal (caracal)

Caracal aurata (African golden cat)

Ocelot lineage
Leopardus

Leopardus pardalis (ocelot)

Leopardus wiedii (margay)

Leopardus jacobita (Andean mountain cat)

Leopardus colocolo (Pampas cat)

Leopardus geoffroyi (Geoffroy's cat)

Leopardus guigna (kodkod)

Leopardus tigrinus (oncilla or tigrina)

Lynx lineage
Lynx

Lynx rufus (bobcat)

Lynx canadensis (Canadian lynx)

Lynx lynx (Eurasian lynx)

Lynx pardinus (Iberian lynx)

Puma lineage
Acinonyx

Acinonyx jubatus (cheetah)

Puma

Puma concolor (cougar)

Herpailurus

Herpailurus yagouaroundi (jaguarundi)

Leopard cat lineage
Otocolobus

Otocolobus manul (Pallas's cat)

Prionailurus

Prionailurus rubiginosus (rusty-spotted cat)

Prionailurus bengalensis (leopard cat)

Prionailurus viverrinus (fishing cat)

Prionailurus planiceps (flat-headed cat)

Felis
 

Felis chaus (jungle cat)

Felis nigripes (black-footed cat)

Felis margarita (sand cat)

Felis bieti (Chinese mountain cat)

Felis lybica (African wildcat)

Felis silvestris (European wildcat)

Felis catus (domestic cat)

Domestic cat lineage    

Habitat and ecology[edit]

Cat species are native to every continent except Australasia and Antarctica. Some are adapted to desert environments, some to wetlands, some to high altitude mountainous terrain. Those cat species living in forests are generally agile climbers. All cat species are obligate carnivores and require meat. Apart from the lion, wild cats are generally solitary and secretive. Feral domestic cats form colonies. Cheetah males are known to live and hunt in groups. Activity pattern of cat species ranges from nocturnal to crepuscular and diurnal, depending on their preferred prey species.[11]

See also[edit]

References[edit]

  1. ^ Wozencraft, W.C. (2005). "Felidae". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–548. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ^ a b McKenna, M. C.; Bell, S. K. (2000). "Family Felidae Fischer de Waldheim, 1817:372. Cats". Classification of Mammals. Columbia University Press. p. 230. ISBN 978-0-231-11013-6.
  3. ^ Salles, L. O. (1992). "Felid phylogenetics: extant taxa and skull morphology (Felidae, Aeluroidea)" (PDF). American Museum Novitates (3047).
  4. ^ Hemmer, H. (1978). "Evolutionary systematics of living Felidae – present status and current problems". Carnivore. 1: 71–79.
  5. ^ Johnson, W. E.; Dratch, P. A.; Martenson, J. S.; O'Brien, S. J. (1996). "Resolution of recent radiations within three evolutionary lineages of Felidae using mitochondrial restriction fragment length polymorphism variation". Journal of Mammalian Evolution. 3 (2): 97–120. doi:10.1007/bf01454358.
  6. ^ Christiansen, P. (2008). "Evolution of skull and mandible shape in cats (Carnivora: Felidae)". PLOS ONE. 3 (7): e2807. Bibcode:2008PLoSO...3.2807C. doi:10.1371/journal.pone.0002807. PMC 2475670. PMID 18665225.
  7. ^ a b Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D. W.; Loveridge, A. J. Biology and Conservation of Wild Felids. Oxford, UK: Oxford University Press. pp. 59–82. ISBN 978-0-19-923445-5.
  8. ^ a b Pocock, R. I. (1917). "The classification of the existing Felidae". Annals and Magazine of Natural History. Series 8. XX (119): 329–350. doi:10.1080/00222931709487018.
  9. ^ a b Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O’Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z.; Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News. Special Issue 11.
  10. ^ a b c d Pocock, R. I. (1917). "VII.—On the external characters of the Felidæ". The Annals and magazine of natural history; zoology, botany, and geology. 8. 19 (109): 113−136.
  11. ^ a b c d e f g h i j k Sunquist, M.; Sunquist, F. (2002). "What is a Cat?". Wild Cats of the World. Chicago: University of Chicago Press. pp. 5–18. ISBN 978-0-226-77999-7.
  12. ^ a b c d e Kitchener, A. C.; Van Valkenburgh, B.; Yamaguchi, N. (2010). "Felid form and function". In Macdonald, D.; Loveridge, A. Biology and Conservation of wild felids. Oxford: Oxford University Press. pp. 83−106.
  13. ^ a b Pocock, R. I. (1939). "Felidae". The fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. London: Taylor and Francis. pp. 191–330.
  14. ^ Ewer, R. F. (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. Retrieved 27 January 2013.
  15. ^ Hewett, J. P.; Hewett Atkinson, L. (1938). Jungle trails in northern India: reminiscences of hunting in India. London: Metheun and Company Limited. Archived from the original on 2017-01-18.
  16. ^ Heptner, V. G.; Sludskij, A. A. (1992) [1972]. "Tiger". Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola [Mammals of the Soviet Union. Volume II, Part 2. Carnivora (Hyaenas and Cats)]. Washington DC: Smithsonian Institution and the National Science Foundation. pp. 95–202.
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