Overview

Comprehensive Description

Sunquist and Sunquist (2009) recognized 37 species in the cat family (Felidae), but noted that this number would likely grow somewhat as further taxonomic studies confirm that some currently recognized species actually are composed of distinct populations that warrant recognition as species. Although the number of recognized species has been relatively stable for a century or two, the recognition of genera has been far more dynamic, with the number generally accepted ranging from just two or three to more than a dozen (Sunquist and Sunquist 2009 recognized 14 genera).

There are native felids across all major regions of the Earth except for Australasia and the polar regions (domestic cats have been introduced to many remote oceanic islands and Australia). They may be found from sea level to 6000 meters and in habitats ranging from deserts to savannahs to tropical rainforests, temperate forests, and boreal forests, but around 90% of felid species are associated with forests and woodlands. Some species have relatively narrow geographic and ecological ranges, but a few are extreme generalists. The Puma (Puma concolor) has a range spanning more than 100 degrees of latitude from the Canadian Yukon to the Straits of Magellan and is found from the moist coniferous forests of British Columbia in Canada to the deserts of the southwestern United States, the tropical forests of Central and South America, and south to the cold, dry grasslands of Patagonia. Leopards (Panthera pardus) have a similarly broad distribution in the Old World, ranging from South Africa to the Russian Far East and occurring in habitats from desert to tropical forest. The Colocolo (Leopardus colocolo) and Leopard Cat (Prionailurus bengalensis) (of South America and Asia, respectively) are also notable habitat generalists, although in the case of the Colocolo what is currently treated as a single species may in fact be several cryptic species.The Leopard Cat has the broadest distribution of all small Asian felids, occurring from southern India to the islands of the Sunda Shelf and north to the Russian Far East and ranging across diverse habitats from sea level to 3000 meters in the Himalayas.

The extant felids are relatively homogeneous in their morphology, so much so that the skeletons of two species as different as a Lion (Panthera leo) and a Tiger (Panthera tigris) would be difficult for a non-specialist to distinguish. A typical felid has a rounded head, a relatively flat face, facial whiskers, large eyes and ears, and a sleek and streamlined body with muscular legs. That said, the felids as a group are remarkably variable in size, probably more so than any other mammal family, ranging acoss two orders of magnitude in mass from the 2 to 3 kilogram Black-footed Cats (Felis nigripes), Kodkods (Leopardus guigna), and Rusty-spotted Cats (Prionailurus rubiginosus) to 300 kilogram Tigers. In most species, males tend to be around 5 to 10% larger than females. Many felid species with large geographic ranges exhibit size variation consistent with Bergmann's Rule. For example, Pumas from low latitudes (i.e., closer to the equator) have skulls that average as much as 25% shorter than higher latitude Pumas and an even greater effect of latitude is seen on body weight.

Most felids are nocturnal and nearly all felids are solitary as adults (with only Lions being truly social, although Cheetahs [Acinonyx jubatus] are also somewhat social). Both vocalizations and visual signals are used extensively in communication. Felids are believed to have a relatively poor sense of smell relative to other carnivores, but use scent-marking extensively to communicate with conspecifics.

Felids are sometimes referred to as hypercarnivores because of the much higher proportion of protein they require in their diet, much more than most other mammals. The largest felids are predators of very large mammals. The dominant large cats are the Lion in Africa, the Tiger in Asia, and the Jaguar (Panthera onca) in South America. The medium-sized cats such as the Puma, Snow Leopard (Panthera uncia), and Leopard are able to kill prey their own size, but much of their diet consists of smaller prey between 2 and 40 kilograms and their diets tend to include a much larger range of species than do the diets of the big cats, which tend to feed on just a few species in any particular ecosystem. Small felids, such as Ocelots (Leopardus pardalis), Bobcats (Lynx rufus), Black-footed Cats, and Jungle Cats (Felis chaus), also feed on mammals, but their diets frequently include birds, reptiles, amphibians, and insects as well, with most prey items less than one kilogram. A few small cats are relatively specialized predators, e.g., the Fishing Cat (Prionailurus viverrinus), which captures fishes and frogs by wading in shallow water or waiting on the bank, often plunging its head completely underwater to seize a fish. Fishing Cats have reportedly been seen swimming underwater to catch coots or ducks. Servals (Leptailurus serval) are specialized on small mammals and Canadian Lynx (Lynx canadensis) feed heavily on Snowshoe Hares (Lepus americanus).

Coat pattern is highly variable both among and within felid species.  An especially striking example of intraspecific variation is the melanism that has been recorded for a number of felid species living in tropical humid and densely vegetated habitats. Best known are the "black panthers", which are melanistic Leopards (in the Old World) or Jaguars (in the New World).

Many species of felids have declined dramatically over the past century or two due to human impacts, with declining ranges and shrinking populations resulting from habitat loss, declining prey populations, and direct persecution for trade, predator control, and sport.  Some felid species are believed to be naturally rare (e.g., the extremely poorly known Bay Cat [Catopuma badia] of Borneo and the three smallest felids, the Kodkod of Chile and Argentina, the Rusty-spotted Cat of India and Sri Lanka, and the Black-footed Cat of South Africa), making them more vulnerable to new threats. Sunquist and Sunquist (2009) provided an overview of the conservation threats facing many of the world's felids as of 2009.

(Sunquist and Sunquist 2009 and references therein)

  • Eizirik, E., W.J. Murphy, K.P. Koepfli, W.E. Johnson, J.W. Dragoo, and S.J. O'Brien. 2010. Pattern and timing of the diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences. Molecular Phylogenetics and Evolution 56: 49-63.
  • Johnson, W.E., E. Eizirik, J. Pecon-Slattery, W.J. Murphy, A. Antunes, E. Teeling, and S.J. O'Brien. 2006. The late miocene radiation of modern felidae: A genetic assessment. Science 311: 73-77.
  • Sunquist, M.E. and F.C. Sunquist. 2009. Family Felidae (Cats). Pp. 54-168 in: Wilson, D.E. & Mittermeier, R.A., eds. Handbook of the Mammals of the World. Volume 1. Carnivores. Lynx Edicions, Barcelona.
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Diversity

With the exception of Antarctica, Australia, New Zealand, Madagascar, Japan, and most oceanic islands, native populations of cats are found worldwide, and one species, domestic cats, have been introduced nearly everywhere humans currently exist. Although some authorities recognize only a few genera, most accounts of Felidae recognize 18 genera and 36 species. With the exception of the largest cats, most are adept climbers, and many are skilled swimmers. Most felids are solitary. Often, felids are separated into two distinct subgroups, large cats and small cats. Generally, small cats are those that, due to a hardening of the hyoid bone, have an inability to roar. Felidae consists of 2 subfamilies, Pantherinae (e.g., lions and tigers) and Felinae (e.g., bobcats, pumas, and cheetahs).

Felids are perhaps the most morphologically specialized hunters of all carnivores, often taking prey as large as themselves and occasionally taking prey several times their own size. Unlike other carnivores, felids rely almost exclusively on prey that they have killed themselves. They are agile hunters, hunting mostly at night, with diets consisting of fresh meat or carrion. Felids are found in all terrestrial habitats except treeless tundra and polar ice caps.

The first cat-like mammals appeared around 60 million years ago (MYA) during the Eocene and culminated in the most specialized of the saber-tooths, Barbourofelis fricki. However, the phylogeny of saber-tooths and their ancestors (Nimravidae) is the subject of considerable debate and fossil evidence for these cat-like mammals does not exist after the Miocene. True felids first appeared during the early Oligocene and, although early ancestors of present day felids had short upper canines, felid radiations that occurred during the Miocene and Pliocene, such as Smilodon, appeared to specialize on large herbivores and had large, saber-like upper canines. Early felids were divided into two subfamilies, Machairodontinae (saber-toothed cats) and Felinae (conical-toothed cats). The many genera of saber-toothed cats are divided into three tribes (Metailurini, Homotheriini, and Smilodontini). Living and extinct conical-toothed cats are placed in one subfamily and one tribe, the Felini, but controversy surrounds generic-level classification of felids. Modern cats are closely related to hyenas, mongooses, and civets. These families, including the families Eupleridae and Nandiniidae, are in the suborder Feliformia.

  • International Union for Conservation of Nature and Natural Resources. 2008. "2008 IUCN Red List of Threatened Species" (On-line). Accessed March 26, 2009 at http://www.iucnredlist.org/search.
  • Boorer, M. 1970. Wild Cats. New York: Grosset & Dunlap.
  • Clutton-Brock, J., D. Wilson. 2001. Cats. Pp. 208-215 in D Burnie, D Wilson, eds. Smithsonian Institution: Animals. New York, NY: DK Publishing.
  • Colby, C. 1964. Wild Cats. New York: Duell, Sloan, and Pearce.
  • Denis, A. 1964. Cats of the World. Boston: The Riverside Press Cambridge.
  • Grzimek, B. 2003. Cats (Felidae). Pp. 369-392 in H Michael, K Devra, G Valerius, M Melissa, eds. Family: Felidae, Vol. 14, 2 Edition. Farmington Hills, Michigan: The Gale Group.
  • Kelsey-Wood, D. 1989. The Atlas of Cats of the World: Domesticated and Wild. Neptune City, NJ: T.F.H. Publications.
  • O'Brian, S. 2001. Cat Family. Pp. 8-39 in D Macdonald, ed. The Encyclopedia of Mammals. Oxford, UK: Andromeda Oxford Limited.
  • Turner, A. 1997. The Big Cats and Their Fossil Relatives: an Illustrated Guide to Their Evolution and Natural History. New York: Columbia University Press.
  • UCMP, 2010. "Geologic Time Scale" (On-line). University of California Museum of Paleontology (UCMP). Accessed March 20, 2011 at http://www.ucmp.berkeley.edu/education/explorations/tours/geotime/guide/geologictimescale.html.
  • Vaughan, T., J. Ryan, N. Czaplewski. 2000. Mammalogy. Fourth Edition. Philadelphia, PA: Brooks/Cole-Thomson Learning.
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Description of Felidae

The cats. Felids are the most strictly carnivorous of the sixteen mammal families in the order Carnivora. The most familiar felid is the domestic cat, which first became associated with humans about 10,000 years ago, but the family includes all other wild cats including the big cats.  Discounting extinct species, the living felids belong to either the Pantherinae (lion, tiger, jaguar, and leopard), and Felinae (cougar, cheetah, lynxes caracal, and domestic cat).  The first felids emerged dabout 25 million years ago. In prehistoric times, there was a third group called the Machairodontinae, which included the saber-toothed cats such as the well known Smilodon.   There are 40 known species of felids in the world today. They originated in Asia and spread across continents by crossing land bridges. 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 Isthmus of Panama. The Panthera species are the oldest and the Felis species are the youngest.   Felids are purely carnivorous animals, subsisting almost entirely on other vertebrates. Aside from the lion, they are solitary. Most are secretive animals, often nocturnal, and live in relatively inaccessible habitats. Around three-quarters of cat species live in forested terrain, and they are generally agile climbers. However, felids may be found in almost any environment, with some species being native to mountainous terrain or deserts. Wild felids are native to every continent except Australia and Antarctica.  The various species of felid vary greatly in size. One of the smallest is the Black-footed Cat at between 35-40 cm in length, while the largest and most powerful is the Siberian tiger. Compared with many other mammals, they have relatively short faces, and good binocular vision.  The fur of felids takes many different forms, being much thicker in those species that live in cold environments, such as the Snow Leopard. The colour of felids is also highly variable, although brown to golden fur is common in most species, often marked with distinctive spots, stripes, or rosettes. Many species also have a \"tear stripe,\" a black stripe running from the corner of each eye down the side of the nose.  The tongue of felids is covered with horny papillae, which help to rasp meat from their prey. Almost all felids have fully retractable claws (one exception is the cheetah). Cats have five toes on their forefeet and four on their hindfeet, reflecting their reliance on griping and holding down their pray with their claws.  Felids have relatively large eyes, 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 cat eyes their distinctive shine.  The ears of felids are also large, and in the smaller cats especially sensitive to high-frequency sounds. Felids have a highly developed sense of smell, although not as much so as 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.  Felids possess highly sensitive vibrissa (whiskers) set deep within the skin, and provide the cat with sensory information about the slightest air movement around it. For this reason they are very helpful for a nocturnal hunter. Most felids are able to land on their feet after a fall, an ability which relies on vision and the sense of balance acting together.  The canine teeth are large, reaching exceptional size in the extinct saber-tooth species. The upper third premolar and lower molar are adapted as carnassial teeth, suited to tearing and cutting flesh.  The jaws of felids can only move vertically. This prevents them from being able to chew, but makes it easier for their powerful masseter jaw muscles to hold struggling prey.   Text after Wikipedia, 3rd August 2008.
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Distribution

Geographic Range

Felids are native to every continent except Australia and Antarctica. Excluding domestic and feral cats (Felis catus), which are globally distributed, felids can be found everywhere except Australia, New Zealand, Japan, Madagascar, polar regions, and many isolated oceanic islands.

Biogeographic Regions: nearctic (Introduced , Native ); palearctic (Introduced , Native ); oriental (Introduced , Native ); ethiopian (Introduced , Native ); neotropical (Introduced , Native ); australian (Introduced ); oceanic islands (Introduced )

Other Geographic Terms: holarctic ; cosmopolitan

  • Feldhamer, G., L. Drickamer, S. Vessey, J. Merritt. 1999. Mammalogy: Adaptation, Diversity, and Ecology. Second Edition. Boston, MA: McGraw Hill.
  • Nowell, K., P. Jackson. 1996. Wild Cats. Cambridge, U.K.: International Union for Conservation of Nature.
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Physical Description

Morphology

Physical Description

All felids bear a strong resemblance to one another. Unlike members of the family Canidae, felids have a short rostrum and tooth row, which increases bite force. Loss or reduction of cheek teeth is particularly apparent in felids, which have a typical dental formula of 3/3, 1/1, 3/2, 1/1 = 30. In most species, the upper premolar is significantly reduced and in Lynx, has been completely lost. Felids have well developed carnassials. Their cheek teeth are secodont and are specialized for shearing. Felid canines tend to be long and conical and are ideal for puncturing prey tissues with minimal force. Besides having a short rostrum, felids also have large bullae that are divided by a septum; no alisphenoid canal, and paroccipital processes flattened against the bullae. Felids also have a vestigial or absent baculum and retractable claws. Distal segments of digits in the relaxed position are pulled back and up into a sheath by an elastic segment, which prevents claws from becoming blunt. Cheetahs are the exception as they cannot retract their claws and, when attacking prey, they tend to run into them so that they fall, much like canids. Cats have five toes on their forefeet and four on their hindfeet. They are digitigrade, and their metapodials are moderately long but never fused.

Felids range in body mass from 2 kg in black-footed cats (Felis negripes) to 300 kg in tigers (Panthera tigris), and exhibit sexual dimorphism, with males being larger and more muscular than females. In some species, such as lions (Panthera leo), males may also have ornamentation that is used to attract potential mates. Throughout their range, felid coats are longest where temperatures tend to be coldest (e.g., snow leopards). Felids exhibit a wide range of colors, from black to orange to white, and many species have cryptically colored coats containing rosettes, spots, and stripes that help camouflage them while hunting for prey. While melanistic variants (solid black) are common in many species, completely white individuals tend to be rare. A great deal of color variation can occur within individual species and newborns tend to have different coloration than adults. For example, adult cougars (Puma concolor) rarely have spots while kittens almost always have spots. In general, the ventral surface of felids tends to be pale while the face, tail, and back of the ears often have black or white markings.

Felids have a number of morphological adaptations that have allowed them to become the most adept hunters in the order Carnivora. They have digitigrade posture that results in a rapid stride rate and powerful forelimbs that help them capture and retain large prey. Often, felids are cryptically colored, which helps camouflage them while hunting. In addition, most felids have large eyes and exceptional vision. In nocturnal species, the tapetum lucidum helps intensify limited light. Many species also have large semi-rotating ears. Finally, the felid tongue has a sandpaper-like texture due to posteriorly directed papillae on its dorsal surface, which are thought to help retain food in the mouth and remove tissue from the bones of prey.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger; ornamentation

  • Kitchener, A. 1991. The Natural History of the Wild Cats. Ithaca, New York: Comstock Publishing Associates.
  • Walker, E. 1975. Mammals of the World. Baltimore and London: The John Hopkins University Press.
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Ecology

Habitat

Cats are found in all terrestrial habitats except treeless tundra and polar ice regions. Most species are habitat generalists and can be found in a wide range of environments. However, few have adapted to a limited range of habitats. For example, optimal habitat for sand cats (Felis margarita) consists of sandy and stony deserts. Domestic and feral cats (F. catus) are ubiquitous globally and are especially pervasive in urban and suburban areas.

Habitat Regions: temperate ; tropical ; terrestrial

Terrestrial Biomes: taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp ; bog

Other Habitat Features: urban ; suburban ; agricultural ; riparian

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Trophic Strategy

Food Habits

Morphologically, felids are considered the most specialized of all carnivores in the order Carnivora. They are at top of the food web in most ecosystems, as their diet consists almost entirely of animals. Occasionally, felids ingest grass to help pass fur balls, a by-product of constant grooming. Some genera ingest fruit to help offset water requirements. Felids may eat the viscera (i.e., internal organs) of prey, thus consuming partially digested plant biomass. Although they typically hunt for large prey (e.g., Perissodactyla and Artiodactyla), when the opportunity arises large cats may eat carrion as well. Small cats predominantly prey upon rodents and rabbits or hares. When available, small cats also feed upon reptiles, amphibians, birds, fish, crustaceans, and arthropods. Some species cache food and may drag prey carcasses into nearby trees prior to feeding (e.g., Panthera pardus). Fishing cats and flat-headed cats are unique among felids, as they are especially adapted for preying upon fish and frogs.

Foraging Behavior: stores or caches food

Primary Diet: carnivore (Eats terrestrial vertebrates, Piscivore , Scavenger )

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Associations

Ecosystem Roles

Felids are apex predators that initiate top-down control and are often considered keystone species in their native habitats. Often preying upon the most vulnerable of individuals (e.g., young, old, or ill), felids promote robust prey populations that exhibit decreased vulnerability to disease and prevent overgrazing by large herbivores. For example, evidence suggests that white-tailed deer in Bear Island, Florida avoid forest habitat based on the presence or absence of Florida panthers. However, bobcats, which typically prey upon small mammals, opportunistically prey on deer in open habitat. Thus, in their attempt to avoid one felid predator, white-tailed deer have become increasingly vulnerable to another.

Domestic and wild felids are vulnerable to a large number of endoparasites including flatworms (Platyhelminthes), roundworms (Nematoda), thorny-headed worms (Acanthocephala), tongue worms (Pentastomida), and parasitic protozoa (Apicomplexa). Common felid ectoparasites consist of ticks and mites (Acari), sucking lice (Phthiraptera), fleas (Siphonaptera), mosquitoes (Culicidae), and flies (Diptera).

Ecosystem Impact: keystone species

Commensal/Parasitic Species:

  • Maehr, D., M. Orlando, J. Cox. 2005. Large carnivores, herbivores, and omnivores in South Florida: an evolutionary approach to conserving landscape and biodiversity. Pp. 293-313 in J Ray, K Redford, R Steneck, J Berger, eds. Large Carnivores and the Conservation of Biodiversity. Washington D. C.: Island Press.
  • Millan, J., F. Ruiz-Fons, F. Marquez, M. Viota, J. Lopez-Bao, M. Martin-Mateo. 2007. Ectoparasites of the endangered Iberian lynx (Lynx pardinus) and sympatric wild and domestic carnivores in Spain. Medical and Veterinary Entomology, 21: 248-254.
  • Patton, S., A. Rabinowitz. 1994. Parasites of wild Felidae in Thailand: a coprological survey. Journal of Wildlife Diseases, 30/3: 472-475.
  • Patton, S., A. Rabinowitz, S. Randolph, S. Johnson. 1986. A coprological survey of parasites of wold neotropical felidae. The Journal of Parasitology, 72/4: 517-520.
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Predation

Felids are typically apex predators (i.e., predators with no predators of their own), but young are vulnerable to predation until they are capable of defending themselves. Many species are cryptically colored, which allows them to remain camouflaged while in their native habitat. Although not an act of predation, many large cats are intolerant of heterospecific felids. For example, lions readily kill leopards, which are known to kill cheetahs. During attempted pride takeovers, male lions commit infanticide as a way of inducing estrus in pride females and eliminating the offspring of rival males. About one quarter of lion cub deaths can be attributed to infanticide, which also occurs in pumas.

Anti-predator Adaptations: cryptic

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Known prey organisms

  • A. C. Twomey, The bird population of an elm-maple forest with special reference to aspection, territorialism, and coactions, Ecol. Monogr. 15(2):175-205, from p. 202 (1945).
  • Myers, P., R. Espinosa, C. S. Parr, T. Jones, G. S. Hammond, and T. A. Dewey. 2006. The Animal Diversity Web (online). Accessed February 16, 2011 at http://animaldiversity.org. http://www.animaldiversity.org
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Life History and Behavior

Behavior

Communication and Perception

Felids have acute senses of smell, hearing, and sight. In addition to the tapetum lucidum, a layer of reflective tissue in the eye of many vertebrates, felids have a modified pupil that allows for excellent vision in a wide range of environments. The felid pupil consists of a vertical slit that expands in low light conditions and contracts in high light conditions. Felids have relatively large pinnae that can rotate to allow for multidirectional hearing without rotating their head. Well-developed vibrissae, which are located above the eyes, on the muzzle, and on the ventral surface of forepaws between the digits, play an important role in tactile sensory reception. Similar to other carnivores, felids have haptic receptors inside their digits that allows them to sense temperature, pressure, and other stimuli.

Felids are solitary animals that scent mark territories with facial glands and urine. They also mark territorial boundaries by clawing tree trunks. Like many vertebrates, felids have a vomeronasal organ, or Jacobson's organ, that allows them to detect pheromones. This olfactory sense organ is found at the base of the nasal cavity and plays an important role in conspecific interactions, especially those related to reproduction. For example, after smelling the genital area or urine of a potential mate, males curl their upper lip toward their nostrils (i.e., the Flehmen response). Using the vomeronasal organ, this allows males to assess the mating condition and quality of potential mates. It is thought that input from the vomeronasal organ and the olfactory bulbs significantly contribute to mating activity.

Due to their nocturnal and solitary lifestyles, investigating audible communication in felids has proven difficult. However, the calls of many carnivores are known to signal individual recognition and territorial boundaries. It is thought that by observing domestic cats (Felis catus), one can hear a majority of the sounds made by most felids. They purr, meow, growl, hiss, spit, and scream. The hyoid apparatus of small-bodied cats is hardened, resulting in an inability to roar. Large-bodied cats have the capability to roar, which is thought to serve as a form of long-distance communication. For example, lions typically roar at night to advertise territories. Research suggests that lionesses can identify the sex of a roaring individual and lionesses respond differently to different numbers of roaring individuals.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: pheromones ; scent marks

Perception Channels: visual ; acoustic

  • Packer, C. 2001. Why lions roar: long distance vocal communication in African prides. Pp. 16-17 in D Macdonald, ed. The Encyclopedia of Mammals. Oxford, UK: Andromeda Oxford Limited.
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Life Expectancy

Lifespan/Longevity

Although many cats do not live beyond their first birthday, felid lifespan ranges from 15 to 30 years old. In the wild, juvenile deaths are normally due to predation. In captivity, however, juvenile deaths are often due to stillbirths, cannibalism, maternal neglect, hypothermia, and congenital disorder.

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Reproduction

Felids are most often classified as polygynous but can exhibit polygynandrous behavior as well. Estrous lasts from 1 to 21 days and females may have multiple estrous cycles until they become pregnant. Females advertise estrus to potential mates through vocalizations, scent marking, and restlessness. As with most polygynous species, males compete for access to mates via displays and fighting, and successful males court mates through vocalizations and direct physical contact (e.g., rubbing on the female). During courtship, successful males may also approach receptive females with their head lowered. While the act of copulation lasts less than a minute, multiple copulations can occur over a period of several days, which may help induce ovulation. After several days, males may leave in order to find additional estrous females, in which case another male takes his place.

In felids, male territories often encompass those of multiple females (for an exception see Panther leo) and males mate with females that reside within his territory. Most conspecific interactions occur during mating season or as a result of territorial disputes among rival males. Indirect interactions via scent markings or vocalizations help reduce the number of fatal interactions.

Mating System: polygynous ; polygynandrous (promiscuous)

The act of copulation is aggressive and brief and may be repeated multiple times an hour for several days. Repeated copulation is thought to induce ovulation in females. Most species are polygynous and polyestrous, with estrous cycles lasting from from 1 to 3 days. Most felids are non-seasonal breeders, but in areas of extreme climatic or prey variability, parturition occurs during the most favorable times of the year. Small-bodied cats tend to have 3 litters per year, while large cats average 1 litter every 18 months. The interval between birthing events may depend on maturation rates of young, body size, food availability, or recent loss of litter. For example, if a female loses her litter, she can come into estrus within a few weeks. Although most litters contain 2 to 4 cubs, females can give birth to as many as 8 cubs in a litter. Gestation lasts from 2 months in small cats to 3 months in lions and tigers.

Felid cubs are born altricial, as newborns are often blind and deaf, rendering them defenseless. Mothers often hide newborns in dens, rock crevices, or tree hollows until they are mobile. Cubs remain with their mother until they can hunt on their own. Weaning begins at the introduction of solid food and ranges in length from 28 days (domestic cats) to 100 days (lions and tigers). Felids reach sexual maturity in less than a year for small cats and up to 2 years for large cats. Typically, cats do not produce their first litter until they have established a home range, which usually does not occur until they are 3 or 4 years of age. Although age of independence is highly variable, many species become independent around 18 months of age. Unlike most felids, lions are very social and females take turns nursing young born to other pride members (i.e., communal nursing) while absent mothers are hunting for food.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); induced ovulation ; viviparous

With the exception of lions, females are the sole caretakers of young felids. Mothers hide their cubs in dens, rock crevices, or tree hollows while they are away hunting and young hide until she returns. Weaning begins at the introduction of solid food, around 28 days in domestic cats (Felis domesticus) and 100 days in lions. Females teach cubs how to stalk, pounce, and kill. Weaning is complete when cubs can eat meat and help hunt for prey. Juvenile felids spend a majority of their time “role playing,” which helps develop important hunting skills. Juveniles are independent once they become competent hunters, though they may remain in their mother’s territory for up to a year before they establish their own. Most felids do not begin reproducing until they have their own territories. Although male lions use infanticide to eliminate unrelated young during pride takeover events, they also provide a significant degree of parental care to their own offspring, protecting cubs while they feed and allowing mothers to rest.

Parental Investment: altricial ; female parental care ; pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); extended period of juvenile learning

  • Grzimek, B. 2003. Cats (Felidae). Pp. 369-392 in H Michael, K Devra, G Valerius, M Melissa, eds. Family: Felidae, Vol. 14, 2 Edition. Farmington Hills, Michigan: The Gale Group.
  • Kitchener, A. 1991. The Natural History of the Wild Cats. Ithaca, New York: Comstock Publishing Associates.
  • Patterson, B. 2007. On the Nature and Significance of Variability in Lions (Panthera leo). Evolutionary Biology, 34: 55-60.
  • Ramel, G. 2008. "The Cats (Family Felidae; Order Carnivora)" (On-line). The Cats. Accessed February 24, 2009 at http://www.earthlife.net/mammals/cats.html.
  • Sunquist, M., F. Sunquist. 2002. Wild Cats of the World. Chicago and London: The University of Chicago Press.
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Evolution and Systematics

Functional Adaptations

Functional adaptation

Skeletal construction provides shock absorption: cats
 

The skeleton of a cat allows it to absorb shocks to its forelimbs because it has no direct skeletal connection between its collarbone and vertebral column.

     
  "Cats have no direct skeletal connection through a collarbone between the bones of their forelimbs (pectoral girdle) and those of their vertebral columns. In effect, they have shock-mounted forelimbs, which cushion a landing after a jump. None of these schemes reduce the extremes of velocity one bit; what they reduce are the velocity gradients." (Vogel 2003:77)
  Learn more about this functional adaptation.
  • Steven Vogel. 2003. Comparative Biomechanics: Life's Physical World. Princeton: Princeton University Press. 580 p.
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Functional adaptation

Olfactory bulb detects X-rays: cat
 

The sensory system of cats detects X-ray radiation with the olfactory bulb, rather than the eyes.

   
  "In 1965, a team of biologists at the Veterans Administration Hospital in Long Beach, California, performed experiments that seemed to show that cats could detect X rays. In conditioning experiments, cats reacted to five-second exposures of X-ray radiation in order to avoid a mild rebuff. In attempting to pinpoint the body region responsible for this remarkable feat, the researchers found that the olfactory bulb behind the nasal and oral passages was the most responsive region, rather than the eyes." (Shuker 2001:21)
  Learn more about this functional adaptation.
  • Shuker, KPN. 2001. The Hidden Powers of Animals: Uncovering the Secrets of Nature. London: Marshall Editions Ltd. 240 p.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records: 1592
Specimens with Sequences: 3206
Specimens with Barcodes: 260
Species: 40
Species With Barcodes: 40
Public Records: 207
Public Species: 33
Public BINs: 19
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Conservation

Conservation Status

Major challenges to felid populations include habitat loss or fragmentation, management of cat-human interactions, the collection and killing of felids for the pet and fashion trades, and disappearance of natural prey. Additionally, reduced population sizes increase vulnerability to extinction due to natural disasters, epidemics, and inbreeding depression. According to the IUCN Redlist of Threatened Species, 29 of the 36 recognized species of felids are currently in decline, and 5 of the remaining 7 species have insufficient population data to determine demographic trends. Iberian lynx (Lynx pardinus) are listed as critically endangered and are one of the most endangered animals on the planet, with a maximum of 143 individuals remaining in 2 separate breeding populations. Including tigers (Panthera tigris) and snow leopards (Panthera uncia), 6 other species are listed as endangered. CITES, which was created in part over concerns that international fur trade would drive many felids to extinction, currently lists 23 species or subspecies under Appendix I, with all remaining species placed under Appendix II. The North American Endangered Species Act lists 8 species or subspecies of North American felids as threatened or endangered, including jaguars (Panthera onca), ocelots (Leopardus pardalis), and panthers (Puma concolor).

Currently, conservation efforts are focused on habitat preservation, captive breeding, and reintroductions. Numerous cat species have been reintroduced or translocated throughout parts of their range where they were once extinct. Aside from the reintroduction of European wild cats in Bavaria, Canada lynx in northern New York State, and bobcats to Cumberland Island, Georgia, few reintroductions have been truly successful. The majority of felid reintroductions fail due to a lack of careful planning and execution, which is directly linked to a lack of time and money. In addition, a majority of large cat reintroductions fail because management teams don't take into consideration four important points. First, reintroduction efforts must consider the conditions under which past translocation events were successful, especially the movement of animals into established populations. Second, management teams often fail to appropriately train captive-bred animals to be successful predators in their native habitat. Third, prior to a reintroduction or translocation event it is imperative that the various genetic and morphological differences between different subpopulations are well understood. Finally, the support and receptivity of local human communities must be assessed prior to reintroducing a potentially dangerous predator. Many felid populations are currently in decline largely because of persecution by humans. If felid reintroduction is not supported by local communities, such attempts are likely to fail.

In 1996, the IUCN published an action plan for the conservation of large cats, which included a list of 105 "priority projects". The "general conservation plan" called for a number actions that were believed to aid in the conservation of all felid species. For example, the establishment of a "cat conservation center" would result in a centralized data management center that would solicit potential donors for funding and help carry out the directives suggested by the conservation action plan as a whole. In addition to a generalized action plan, species specific action plans were formulated for 43 different cat species. Since 1996, the IUCN's Cat Specialist Group has helped launch numerous research efforts aimed at addressing the conservation goals outlined in their 1996 conservation plan. In 2004, the Cat Specialist Group established a "digital cat library" that contains more than 6,000 "papers and reports relevant to the conservation of wild cats", and in 2005 the first captive bred Iberian lynx litter was born, which served as a giant symbolic leap in the long journey of felid conservation.

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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Where feral domestic cats have been introduced, the diversity of small vertebrates (such as birds, lizards, and small mammals) has significantly declined. Felids attack and kill livestock, which can result in losses for farmers. Wild cats are capable of transmitting pathogens to domestic cats. Large cats occasionally kill and eat people, though a majority of attacks are often the result of accidental confrontations or involve sick or injured animals. In the Sunderbans of India, the largest contiguous parcel of halophytic forest in the world, tigers (Panthera tigris) kill several dozen people each year.

Negative Impacts: injures humans (bites or stings); causes or carries domestic animal disease

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Economic Importance for Humans: Positive

Felids were first domesticated in Egypt between 4,000 and 7,000 years ago. Historically, cat pelts served as a symbol of status and power, a trend that continues to this day. In Africa, felids are often hunted for sport (i.e., trophies) and retaliatory killings by livestock farmers are not uncommon. In addition to their pelts, felids are desired for their claws and teeth. Traditional medicines may incorporate felid by-products, although their efficacy is unproven. Although international trade of wild felids and their by-products is illegal, domestic trade continues in some countries. In the ecotourism industry large cats have significant economic value in Africa and India and are sought out by tourists on both national and private reserves. Small cats primarily prey on rodents, hares, and rabbits, which helps control pest populations throughout much of their range. Large cats commonly prey on large herbivores, which reduces competition between livestock and native ungulates.

Positive Impacts: pet trade ; body parts are source of valuable material; ecotourism ; controls pest population

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Wikipedia

Felidae

Felidae is the biological family of the cats; a member of this family is called a felid. The most familiar felid is the domestic cat, which first became associated with humans about 10,000 years ago, but the family includes all other wild cats, including the big cats.

Extant felids belong to one of two subfamilies: Pantherinae (which includes the tiger, the lion, the jaguar, and the leopard), and Felinae (which includes the cougar, the cheetah, the lynxes, the ocelot, and the domestic cat).

The first felids emerged during the Oligocene, about 25 million years ago. In prehistoric times, a third subfamily, the Machairodontinae, included the "saber-toothed cats", such as the well-known Smilodon. Other superficially cat-like mammals, such as the marsupial sabertooth Thylacosmilus or the Nimravidae are not included in Felidae despite superficial similarities.

Felids are the strictest carnivores of the 13 terrestrial families in the order Carnivora, although the three families of marine mammals comprising the superfamily Pinnipedia are as carnivorous as the felids. Felids are sometimes referred to as hypercarnivores because of the much higher proportion of protein they require in their diet, much more than most other mammals.[3]

Evolution[edit]

The 41 known cat species in the world today all descended from the same ancestor.[1] Cats originated in Asia and spread across continents by crossing land bridges. Testing of mitochondrial and nuclear DNA revealed 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 Panthera genus being the oldest and the Felis genus being the youngest. About 60% of the modern cat species are estimated to have developed within the last million years.[4]

The Felidae's closest relatives are thought to be the Asiatic linsangs.[5] Together with the Viverridae, hyenas, mongooses, and Madagascar carnivores, they form the suborder Feliformia.[6]

Most cat species share a genetic anomaly that prevents them from tasting sweetness.[7]

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.[8] Prior to this discovery, biologists had been largely unable to establish a family tree of cats from fossil records because the fossils of different cat species all look very much alike, differing primarily in size.[citation needed]

Characteristics[edit]

Felids are obligate carnivores, requiring a diet of meat and organs to survive. Aside from the lion, wild felids are generally solitary; feral domestic cats do, however, form colonies. Cheetahs are also known to live and hunt in groups. Felids are generally secretive animals, are often nocturnal, and live in relatively inaccessible habitats. Around three-quarters of cat species live in forested terrain, and they are generally agile climbers. However, felids may be found in almost any environment, with some species being native to mountainous terrain or deserts.

Wild felids are native to every continent except Australasia and Antarctica.

Physical appearance[edit]

Skull of the machairodontine Smilodon (reconstruction)
Lion skull

Felids tend to have lithe and flexible bodies with muscular limbs. In the great majority of species, the tail is between a third and a half the length of the body, although with some exceptions (for example, the bobcat and margay). The limbs are digitigrade with soft toe pads and protractible claws. Compared with most other mammals, the head of cats is highly domed with a short muzzle. The skull possesses wide zygomatic arches and a large sagittal crest, both of which allow for the attachment of strong jaw muscles.[9]

The various species of felids vary greatly in size. One of the smallest is the black-footed cat, measuring 35 to 40 cm (14 to 16 in) long, while the largest in the wild is the tiger, which can grow up to 350 cm (11.5 ft) in length[10] and weigh 300 kilograms (660 lb).

The fur of felids takes many different forms, being much thicker in those species living in cold environments, such as the snow leopard. The color of felids is also highly variable—although brown to golden fur is common in most species—usually marked with distinctive spots, stripes, or rosettes. The only felids to lack significant markings are the lion, puma, caracal, and jaguarundi. Many species exhibit melanism, in which some individuals have an all-black coat.[9]

The tongue of felids is covered with horny papillae, which rasp meat from prey and aid in grooming.

All felids have protractible claws, in other words they have the ability to protract their claws from a retracted, at-rest position. However, in a few species, such as the cheetah, the claws remain visible even when at rest (retracted). The claws are retracted when the animal is relaxed and protracted when they are in use. They are attached to the terminal bone of the toe with a tough ligament; when the animal contracts muscles in the toe to straighten it, the ligament forces the claw outwards.[9] Cats have five toes on their forefeet and four on their hindfeet, reflecting their reliance on gripping and holding down their prey with their claws.[citation needed] In the Felidae, the baculum is shorter than in the Canidae.[11]

Senses[edit]

Comparative illustration of cats of the genus Panthera and Felis by N. N. Kondakov
Feeding postures of Panthera and Felis, as illustrated by N. N. Kondakov

Felids have relatively large eyes, 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 color during the day. However, felids appear to have relatively poor color vision in comparison with humans.[9] This is explained by the fact that felids see moving objects more "colorfully" than still objects, but in an intact environment, are unable to distinguish color tones alone (like turquoise compared to teal, for example).[citation needed]

The external ears of felids are also large, and especially sensitive to high-frequency sounds in the smaller cats. This sensitivity allows them to locate small rodent prey; cats do not apparently produce such sounds.[9]

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. Exceptions include members of the genera Leopardus and Otocolobus.[citation needed]

Felids possess highly sensitive whiskers set deep within the skin, which provide the cat with sensory information about the slightest air movement around it. Whiskers are very helpful to nocturnal hunters.

Most felids are able to land on their feet after a fall, an ability that relies on vision and the sense of balance acting together.[citation needed]

Dentition[edit]

Felids have a relatively small number of teeth compared with other carnivorans, a feature associated with their short muzzles. With a few exceptions, such as the lynx, they have the dental formula: 3.1.3.13.1.2.1. The canine teeth are large, reaching exceptional size in the extinct saber-tooth species. The upper third premolar and lower molar are adapted as carnassial teeth, suited to tearing and cutting flesh.[9]

The jaws of felids can only move vertically. This prevents them from being able to chew efficiently, but makes it easier for their powerful masseter jaw muscles to hold struggling prey.

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; overall, the frequency of felid calls ranges between 50 and 10,000 hertz.

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. Growling is used to indicate a willingness to attack, while the higher-pitched snarl is used when adopting a defensive posture.

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.

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 the cheetah and cougar, and may also be found in the 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, prusten and roaring are found only in big cats. 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.[12] 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.[9] Tigers and jaguars have a very snarly roar, while the roar of leopards and lions is much more throaty.[citation needed]

Social and territorial behavior[edit]

Territorial marking[edit]

Classification[edit]

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

Genetic classification[edit]

Genetic research has provided a basis for a more concise classification for the living members of the cat family based on genotypical groupings.[1][4][13] Specifically, eight genetic lineages have been identified:[14]

The last four lineages are more related to each other than to any of the first four, so form a clade within the Felinae subfamily of family Felidae.

Extant species[edit]

The following is the complete list of genera within family Felidae, grouped according to the traditional phenotypical classification with the corresponding genotypical lineages indicated. It includes all the currently living species of cats.[1]

Fossil felids[edit]

The American lion was one of the abundant Pleistocene megafauna, a wide variety of very large mammals that went extinct about 10,000 years ago.[16]

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 the Machairodontinae, though they count as members of the Felinae in recent times.[17][18] 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.

Genera of the Felidae[edit]

The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997)[2] and Wozencraft (2005) in Wilson and Reeder's Mammal Species of the World for extant genera.[1] Pseudaelurus is included in the Felinae as per McKenna & Bell, despite its basal position in felid evolution. Inconsistent with McKenna and Bell, three additional prehistoric genera, Miracinonyx, Lokontailurus and Xenosmilus, are listed. Sivapanthera is included in the Felinae (not Acinonychinae) and Ischrosmilus is included in the genus Smilodon.

See also[edit]

Cited references[edit]

  1. ^ a b c d e Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 532–548. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ a b c McKenna, Malcolm C.; Susan K. Bell (2000-02-15). Classification of Mammals. Columbia University Press. p. 631. ISBN 978-0-231-11013-6. 
  3. ^ http://eol.org/pages/7674/overview
  4. ^ a b c d e Johnson, W. E., Eizirik, E., Pecon-Slattery, J., Murphy, W. J., Antunes, A., Teeling, E., O'Brien, S. J. (2006). "The late miocene radiation of modern Felidae: a genetic assessment". Science 311 (5757): 73–77. doi:10.1126/science.1122277. PMID 16400146. 
  5. ^ Eizirik E., Murphy W. J., Koepfli K. P., Johnson W. E., Dragoo J. W., O'Brien S. J. (2010). "Pattern and timing of the diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences". Molecular Phylogenetics and Evolution 56 (1): 49–63. doi:10.1016/j.ympev.2010.01.033. PMID 20138220. 
  6. ^ Gaubert P., Veron G. (2003). "Exhaustive sample set among Viverridae reveals the sister-group of felids: the linsangs as a case of extreme morphological convergence within Feliformia". Proceedings of the Royal Society B 270 (1532): 2523–30. doi:10.1098/rspb.2003.2521. PMC 1691530. PMID 14667345. 
  7. ^ Xia, Li; Weihua Li, Hong Wang, Jie Cao, Kenji Maehashi, Liquan Huang, Alexander A. Bachmanov, Danielle R. Reed, Véronique Legrand-Defretin, Gary K. Beauchamp and Joseph G. Brand (July 2005). "Pseudogenization of a Sweet-Receptor Gene Accounts for Cats' Indifference toward Sugar". Public Library of Science 1 (1): 27–35. doi:10.1371/journal.pgen.0010003. PMC 1183522. PMID 16103917. Retrieved 2008-06-30. 
  8. ^ Vella, Carolyn; et al. (2002). Robinson's Genetics for Cat Breeders and Veterinarians, 4th ed. Oxford: Butterworh-Heinemann. ISBN 0-7506-4069-3. 
  9. ^ a b c d e f g Sunquist, Mel; Sunquist, Fiona (2002). Wild cats of the World. Chicago: University of Chicago Press. pp. 5–16. ISBN 0-226-77999-8. 
  10. ^ Vratislav Mazak: Der Tiger. Nachdruck der 3. Auflage von 1983. Westarp Wissenschaften Hohenwarsleben, 2004 ISBN 3-89432-759-6
  11. ^ R. F. Ewer (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. Retrieved 27 January 2013. 
  12. ^ Weissengruber, GE; G Forstenpointner, G Peters, A Kübber-Heiss, and WT Fitch (September 2002). "Hyoid apparatus and pharynx in the lion (Panthera leo), jaguar (Panthera onca), tiger (Panthera tigris), cheetah (Acinonyx jubatus) and the domestic cat (Felis silvestris f. catus)". Journal of Anatomy (Anatomical Society of Great Britain and Ireland) 201 (3): 195–209. doi:10.1046/j.1469-7580.2002.00088.x. PMC 1570911. PMID 12363272. 
  13. ^ O'Brien, S. J., Johnson, W. E. (2005). "Big cat genomics". Annual Review of Genomics and Human Genetics 6: 407–429. doi:10.1146/annurev.genom.6.080604.162151. PMID 16124868. 
  14. ^ Johnson, WE; O'Brien, SJ (1997). "Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes". Journal of molecular evolution. 44 Suppl 1: S98–116. PMID 9071018. 
  15. ^ http://www.bucknell.edu/msw3/browse.asp?id=14000003
  16. ^ Ice Age Animals, Illinois State Museum
  17. ^ van den Hoek Ostende, L. W.; M. Morlo and D. Nagel (July 2006). "Fossils explained 52 Majestic killers: the sabre-toothed cats". Geology Today 22 (4): 150–157. doi:10.1111/j.1365-2451.2006.00572.x. Retrieved 2008-06-30. 
  18. ^ Turner, Alan (1997-04-15). The Big Cats and their fossil relatives. New York: Columbia University Press. p. 60. ISBN 978-0-231-10228-5. 

General references[edit]

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