Overview
Comprehensive Description
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
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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:
- flatworms (Platyhelminthes)
- thorny-headed worms (Acanthocephala)
- roundworms (Nematoda)
- protozoa (Apicomplexa)
- tongue worms (Pentastomida)
- ticks and mites (Acari)
- sucking lice (Phthiraptera)
- fleas (Siphonaptera)
- mosquitoes (Culicidae)
- flies (Diptera)
- 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
Peromyscus maniculatus
Passerina cyanea
Eudocimus ruber
Sarcoramphus papa
Leontopithecus chrysopygus
Callicebus personatus
Neotragus moschatus
Tetracerus quadricornis
Sylvilagus brasiliensis
Based on studies in:
USA: Illinois (Forest)
This list may not be complete but is based on published studies.
- 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
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
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
| Specimen Records: | 1,840 | Public Records: | 185 |
| Specimens with Sequences: | 1,837 | Public Species: | 31 |
| Specimens with Barcodes: | 271 | Public BINs: | 12 |
| Species: | 42 | ||
| Species With Barcodes: | 38 | ||
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Barcode data
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Locations of barcode samples
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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.
- CITES, 2010. "CITES" (On-line). CITES Species Database. Accessed March 29, 2011 at http://www.cites.org/eng/resources/species.html.
- IUCN, 2010. "IUCN SSC/Cat Specialist Group" (On-line). Accessed March 29, 2011 at http://www.catsg.org/catsgportal/20_catsg-website/home/index_en.htm.
- IUCN, 2010. "The IUCN Red List of Threatened Species" (On-line). Mammals. Accessed March 29, 2011 at http://www.iucnredlist.org/initiatives/mammals.
- U. S. Fish and Wildlife Service, 2010. "U. S. Fish and Wildlife Service" (On-line). Endangered Species Program. Accessed March 29, 2011 at http://www.fws.gov/endangered/species/us-species.html.
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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, known as 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.
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Evolution
There are 41 known species of felids in the world today, all of which descended from the same ancestor.[1] This taxon originated in Asia and spread across continents by crossing land bridges. Testing of mitochondrial and nuclear DNA by Warren Johnson and Stephen O'Brien of the US National Cancer Institute demonstrated 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 Isthmus of Panama, with the Panthera genus being the oldest and the Felis genus being the youngest. They estimated 60% of the modern species of cats developed within the last million years.[3]
Most felids 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.[4] Prior to this discovery, biologists had been largely unable to establish a family tree of cats from the fossil record because the fossils of different cat species all look very much alike, differing primarily in size.
The felids' closest relatives are thought to be the Asiatic linsangs,[5] and at one remove the group of civets, hyenas, mongooses, and Madagascar carnivores,[6] with whom they share the Suborder Feliformia. Most felid species share a genetic anomaly that prevents them from tasting sweetness.[7]
Characteristics
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 feral cat 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
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 there are 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.[8]
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 attain up to 350 cm (11.5 ft) in length[9] 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.[8]
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. Although 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.[8] 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 Felidae, the baculum is shorter than in Canidae.[10]
Senses
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.[8] 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 themselves do not apparently produce such sounds.[8]
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. For this reason, 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
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.[8]
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
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 felid 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.[11] 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.[8] 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
Territorial marking
Within the Felidae, male felids usually urinate backwards by curving the tip of the glans penis backward.[12][10] Urine marking by felids is also known as "spray-urinating"[13] or "spray-marking".[14] To identify their territories, male tigers mark trees by spraying urine[15][16] and anal gland secretions, as well as marking trails with scat. Males show a grimacing face, called the Flehmen response, when identifying a female's reproductive condition by sniffing their urine markings.
Lions use urine to mark their territories. They often scrape the ground while urinating, and the urine often flows in short spurts, instead of flowing continuously. They often urinate on vegetation, or on tree trunks at least one meter high.[17]
Male cheetahs mark their territory by urinating on objects that stand out, such as trees, logs, or termite mounds. The whole coalition contributes to the scent. Males will attempt to kill any intruders, and fights result in serious injury or death.[18] When male cheetahs urine-mark their territories, they stand one meter away from a tree or rock surface with the tail raised, pointing the penis either horizontally backward or 60° upward.[19] The odor of cheetah urine (unlike that of other large felids) cannot be easily detected by humans.[20]
Classification
Traditionally, five subfamilies have been distinguished within family Felidae based on phenotypical features: the Felinae, the Pantherinae, the Acinonychinae (cheetahs), the extinct Machairodontinae, and the extinct Proailurinae.[2]
Genetic classification
Genetic research has provided a basis for a more concise classification for the living members of the cat family based on genotypical groupings.[1][21][22] Specifically, eight genetic lineages have been identified:[23]
- Lineage 1 Pantherinae : Panthera, Uncia, Neofelis
- Lineage 2: Pardofelis, Catopuma
- Lineage 3: Leptailurus, Caracal, Profelis
- Lineage 4: Leopardus
- Lineage 5: Lynx
- Lineage 6: Puma, Acinonyx
- Lineage 7: Prionailurus, Otocolobus
- Lineage 8: Felis
The last four lineages are more related to each other than to any of the first four, and so form a clade within the Felinae subfamily of family Felidae.
Extant species
The following is the complete list of genera within family Felidae, grouped according to the traditional phenotypical classification with the corresponding genotypical lineages indicated:
- Family Felidae[1]
- Subfamily Pantherinae
- Subfamily Felinae
- Genus Pardofelis [Lineage 2]
- Genus Catopuma [Lineage 2]
- Genus Leptailurus [Lineage 3]
- Genus Caracal [Lineage 3]
- Genus Profelis [Lineage 3]
- Genus Leopardus [Lineage 4]
- Genus Lynx [Lineage 5]
- Genus Puma [Lineage 6]
- Cougar (Puma concolor)
- Jaguarundi (Puma yagouaroundi)
- Genus Acinonyx[24][Lineage 6]
- Genus Prionailurus [Lineage 7]
- Genus Otocolobus [Lineage 7]
- Genus Felis [Lineage 8]
Fossil felids
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.[26][27] 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
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.
- †Proailurinae
- Felinae
- Pantherinae
- †Machairodontinae
- †Machairodus (Late Miocene, Africa, Eurasia, North America)
- †Homotherium (Pliocene, Pleistocene; Africa, Eurasia, North America)
- †Xenosmilus (Pleistocene; North America)
- †Lokotunjailurus (Latest Miocene; Africa)
- †Miomachairodus (Middle Miocene; Africa, Asia)
- †Hemimachairodus
- †Paramachairodus (Late Miocene; Eurasia, Africa)
- †Megantereon (Pliocene, Pleistocene; North America, Africa, Eurasia)
- †Smilodon (Late Pliocene to Late Pleistocene; North- and South America)
See also
Cited references
- ^ 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.
- ^ 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.
- ^ Mott, Maryann (2006-01-11). "Cats Climb New family Tree". National Geographic News. Retrieved 2006-07-15.
- ^ Vella, Carolyn; et al. (2002). Robinson's Genetics for Cat Breeders and Veterinarians, 4th ed. Oxford: Butterworh-Heinemann. ISBN 0-7506-4069-3.
- ^ 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: 49–63. doi:10.1016/j.ympev.2010.01.033.
- ^ 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, Series B 270 (1532): 2523–30. doi:10.1098/rspb.2003.2521. PMC 1691530. PMID 14667345.
- ^ 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): e3. doi:10.1371/journal.pgen.0010003. PMC 1183522. PMID 16103917. Retrieved 2008-06-30.
- ^ 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.
- ^ Vratislav Mazak: Der Tiger. Nachdruck der 3. Auflage von 1983. Westarp Wissenschaften Hohenwarsleben, 2004 ISBN 3-89432-759-6
- ^ a b R. F. Ewer (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. Retrieved 27 January 2013.
- ^ 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 Indian Armadillo. (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.
- ^ Reena Mathur (2010). Animal Behaviour 3/e. Rastogi Publications. ISBN 978-81-7133-747-7. Retrieved 10 February 2013.
- ^ John W. S. Bradshaw; Rachel A. Casey; Sarah L. Brown (31 January 2013). The Behaviour of the Domestic Cat. CABI. pp. 104–. ISBN 978-1-78064-120-1. Retrieved 22 March 2013.
- ^ Stefan Schulz (17 March 2005). The Chemistry of Pheromones and Other Semiochemicals II. Springer. pp. 249–. ISBN 978-3-540-21308-6. Retrieved 22 March 2013.
- ^ John Seidenstic (1996). Tigers. MBI Publishing Company. pp. 63–. ISBN 978-0-89658-295-8. Retrieved 25 December 2012.
- ^ Burger, B. V.; Viviers, M. Z.; Bekker, J. P. I.; Roux, M.; Fish, N.; Fourie, W. B.; Weibchen, G. (2008). "Chemical Characterization of Territorial Marking Fluid of Male Bengal Tiger, Panthera tigris". Journal of Chemical Ecology 34 (5): 659–671. doi:10.1007/s10886-008-9462-y. PMID 18437496.
- ^ Schaller, George B (2009-10-15). The Serengeti Lion: A Study of Predator-Prey Relations. ISBN 9780226736600.
- ^ Cheetahs (Revised Edition) - Dianne M. MacMillan - Google Books. Books.google.com. Retrieved 2012-11-20.
- ^ T. M. Caro (15 August 1994). Cheetahs of the Serengeti Plains: Group Living in an Asocial Species. University of Chicago Press. pp. 203–. ISBN 978-0-226-09433-5. Retrieved 21 March 2013.
- ^ The Chemistry of Pheromones and Other Semiochemicals II - Google Books. Books.google.com. 2005-03-17. Retrieved 2012-11-20.
- ^ Johnson WE, Eizirik E, Pecon-Slattery J, et al. (January 2006). "The late Miocene radiation of modern Felidae: a genetic assessment". Science 311 (5757): 73–7. doi:10.1126/science.1122277. PMID 16400146.
- ^ O'Brien SJ, Johnson WE (2005). "Big cat genomics". Annu Rev Genomics Hum Genet 6 (1): 407–29. doi:10.1146/annurev.genom.6.080604.162151. PMID 16124868.
- ^ Johnson, W. E.; & O'Brien S. J. (1997). Phylogenetic reconstruction of the Felidae using 16S rRNA and NADH-5 mitochondrial genes. J Mol Evol (1997) 44: S98-116. Retrieved on 2009-11-08 from http://www.ncbi.nlm.nih.gov/pubmed/9071018?dopt=Abstract.
- ^ http://www.bucknell.edu/msw3/browse.asp?id=14000003
- ^ Ice Age Animals, Illinois State Museum
- ^ van den Hoek Ostende, Lars W.; Michael Morlo and Doris 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.
- ^ 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
- Shoemaker, Alan (1996). "1996 Taxonomic and Legal Status of the Felidae". Felid Taxonomic Advisory Group of the American Zoo and Aquarium Association. Archived from the original on 2006-06-12. Retrieved 2006-07-15.
- Turner, A. (1997). The big cats and their fossil relatives. Columbia University Press. ISBN 0-231-10229-1.
- Kirby, G. (1984). "Cat family". In Macdonald, D. The Encyclopedia of Mammals. Facts on File. ISBN 0-87196-871-1.
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