As treated by Wilson and Mittermeier (2009), the mammalian order Carnivora includes 281 species in 128 genera and 16 families, although the number of species and genera recognized can be expected to fluctuate slightly as a result of new research and based on the opinions of different experts (see family taxon pages for more specific information). Of these 16 families, seven are in the suborder Feliformia and nine are in the suborder Caniformia:
Suborder Feliformia: 120 species in 56 genera
Family Nandiniidae (African Palm Civet): 1 species in 1 genus
Family Felidae (cats): 37 species in 14 genera
Family Prionodontidae (linsangs): 2 species in 1 genus
Family Viverridae (civets, genets, oyans): 34 species in 14 genera
Family Hyaenidae (hyenas): 4 species in 4 genera
Family Herpestidae (mongooses): 34 species in 15 genera
Family Eupleridae (Madagascar carnivores): 8 species in 7 genera
Suborder Caniformia: 161 species in 72 genera
Family Canidae (dogs): 35 species in 13 genera
Family Ursidae (bears): 8 species in 5 genera
Family Otariidae (sea lions): 16 species in 7 genera
Family Odobenidae (Walrus): 1 species in 1 genus
Family Phocidae (earless seals): 19 species in 13 genera
Family Ailuridae (red panda): 1 species in 1 genus
Family Procyonidae (raccoons and relatives): 12 species in 6 genera
Family Mephitidae (skunks): 12 species in 4 genera
Family Mustelidae (weasels and relatives): 57 species in 22 genera
Mammals in the order Carnivora are often referred to as carnivores. This can be confusing, however, since in both popular and scientific usage "carnivore" also means simply "meat-eating"--but many mammals not in the order Carnivora also feed largely or exclusively on meat and many members of the Carnivora are quite omnivorous (and African Palm Civets eat mainly fruit!). For clarity, therefore, some authors prefer to use the term "carnivoran" to refer to a member of the order Carnivora.
The order Carnivora is a morphologically and ecologically diverse group that is distributed across the globe and includes many of the animals humans find most "charismatic". Carnivorans range across five or six orders of magnitude in size from the Least Weasel (Mustela nivalis, which can weigh in at as little as 25 g when fully grown; Larivère and Jennings 2009) to the Southern Elephant Seal (Mirounga leonina, weighing in at 2000 to 3000 kg--possibly as much as 5000kg) (Agnarsson et al. 2010; Nyakatura and Bininda-Emonds 2012).
The phylogenetic relationships among the carnivoran families have been the focus of considerable research and controversy, although the structure of some clades is now well established (see references below and EOL family pages for details). One well established finding is that the two suborders comprising the order Carnivora, Feliformia and Caniformia, are indeed reciprocally monophyletic.
- Nyakatura, K. and O.R.P. Bininda-Emonds. 2012. Updating the evolutionary history of Carnivora (Mammalia): a new species-level supertree complete with divergence time estimates. BMC Biology 10:12. doi:10.1186/1741-7007-10-12
- Agnarsson, I., M. Kuntner, and L.J. May-Collado. 2010. Dogs, cats, and kin: A molecular species-level phylogeny of Carnivora. Molecular Phylogenetics and Evolution 54: 726-745.
- 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.
- Flynn, J.J., J.A. Finarelli, S. Zehr, et. al. 2005. Molecular Phylogeny of the Carnivora (Mammalia): Assessing the Impact of Increased Sampling on Resolving Enigmatic Relationships. Systematic Biology 54(2): 317-337.
- Larivère, S. and A.P. Jennings. 2009. Family Mustelidae (Weasels and Relatives). Pp. 564-656 in: Wilson, D.E. & Mittermeier, R.A., eds. Handbook of the Mammals of the World. Volume 1. Carnivores. Lynx Edicions, Barcelona.
- Sato, J.J., M. Wolsan, F.J. Prevosti, et al. 2012. Evolutionary and biogeographic history of weasel-like carnivorans (Musteloidea). Molecular Phylogenetics and Evolution 63: 745-757.
Members of the mammalian order Carnivora are the descendants of a successful late Paleocene radiation of mammals whose primitive food habits were carnivorous. The name "Carnivora" is sometimes taken to mean that members of this group are all carnivorous or that all carnivorous mammals are members of this group. This is not so. Members of Carnivora have diverse food habits, although many are primarily carnivorous, and carnivory is widely distributed in mammals, being found in many other orders including bats, marsupial mammals, primates, and dolphins and whales.
We recognize 13 extant families and around 270 species of Carnivora.
- Stains, H. 1984. Carnivores. Pp. 491-521 in S Anderson, J Jones Jr., eds. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons.
- Vaughan, T., J. Ryan, N. Czaplewski. 2000. Mammalogy, 4th Edition. Toronto: Brooks Cole.
Carnivores are distributed across the world, on all major land masses (except possibly Australia, where the only terrestrial member of Carnivora, dingos (Canis lupus dingo), may have been brought by man) and in all oceans.
Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Introduced ); antarctica (Native ); oceanic islands (Native ); arctic ocean (Native ); indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native ); mediterranean sea (Native )
Other Geographic Terms: cosmopolitan
Most members of the order Carnivora can be recognized by their enlarged fourth upper premolar and first lower molar, which together form an efficient shear for cutting meat and tendon. These teeth are referred to as the carnassial pair. The exceptions are a few forms, such as bears, raccoons, and seals, in which these teeth are secondarily modified.
Besides usually having carnassials, almost all Carnivora retain the primitive number of incisors (3/3); an exception is the sea otter, which has 2/3. The outer (3rd) incisor is often relatively large and canine-like. The canines are large and conical. The number of teeth behind the carnassials varies considerably, from 1/1 in some cats to 2/2 in bears. All teeth are rooted and diphyodont.
The skulls of carnivorans are varied in form. Most have a well-defined, transverse glenoid fossa, and the dominant motion of the jaw is in the dorsal-ventral direction. The primary muscle powering the jaw is the temporal, and sagittal crest associated with the temporal is commonly a conspicuous part of the surface of the skull. Carnivores also have a strong zygomatic arch and a relatively large braincase. The auditory bullae and the turbinals also tend to be large and complex. Carnivores are fairly intelligent animals and most have relatively large brains. All members of Carnivora have simple stomachs.
Pinnipeds are large, perhaps because water conducts heat well and large animals have a low surface area to body mass ratio, which minimizes heat loss due to conduction. Their bodies are insulated by a thick layer of fat called blubber. In all species, the external ears are small or absent, the external genitalia and nipples are hidden in slits or depressions in the body, and the tail is very small. The forelimbs and hindlimbs are transformed into paddles. In both, the proximal limb elements (humerus and femur) remain within the body, and other aspects of the limbs, limb girdles, and spine are highly specialized for swimming. Most species have a relatively short rostrum, and the orbits are large. The cheek teeth are usually homodont (no differentiation along the toothrow), and the teeth are usually shaped like simple cones.
Carnivores tend to be medium-sized animals; too small and they couldn't find enough within their capacity to kill; too large and they wouldn't be able to satisfy their appetites. However, as a group, carnivores span a wide range of body sizes. Least weasels (Mustela nivalis), the smallest carnivores, can weigh as little as 35 grams, and male southern elephant seals (Mirounga leonina), the largest carnivores, can weigh more than 3,600 kg. Many carnivore species are sexually dimorphic in size. Usually males are larger than females (as with fishers, lions, and wolves) but in a few species females are larger than males (as with spotted hyenas). Additionally, males of some species have ornamentation that females lack (as is the case with the inflatable probosci of male elephant seals).
Many carnivores have thick, luxurious coats, though some, like walruses, have coats that are quite sparse. Their fur comes in various colors, including black, white, orange, yellow, red, and almost every imaginable shade of gray and brown. In addition, many carnivores are striped, spotted, blotched, banded, or otherwise boldly patterned. Some species, such as gray wolves, are polymorphic for coat color. Domesticated cats and dogs exhibit thousands of combinations of coat colors and body shapes as a result of selective breeding by humans.
Other Physical Features: endothermic ; heterothermic ; homoiothermic; bilateral symmetry ; polymorphic
Sexual Dimorphism: sexes alike; female larger; male larger; sexes shaped differently; ornamentation
Carnivores occupy just about every type of terrestrial habitat, and many aquatic habitats as well, from the tropics to the poles. They live in forests, deserts, mountains, grasslands, scrublands, tundra, and on open ice. Aquatic and semi-aquatic species live in freshwater rivers, lakes, and marshes, in marine coastal areas, and in the open ocean.
Habitat Regions: temperate ; tropical ; polar ; terrestrial ; saltwater or marine ; freshwater
Terrestrial Biomes: tundra ; taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains ; icecap
Aquatic Biomes: pelagic ; lakes and ponds; rivers and streams; coastal ; brackish water
Wetlands: marsh ; swamp ; bog
Other Habitat Features: urban ; suburban ; agricultural ; riparian ; estuarine
Animal matter makes up a substantial portion of the diet of most carnivorans. However, not all members of Carnivora are carnivorous. Some, such as bears and raccoons, are decidedly omnivorous, and giant pandas are primarily vegetarian. Foods consumed by carnivorans include mammals, birds and eggs, reptiles, amphibians, fish, carrion, insects and other arthropods, earthworms, mollusks, crustaceans, fruit, nuts, tubers, leaves, shoots, and plankton (on which crabeater seals specialize).
Carnivorans have various means of capturing their prey. Some ambush prey, overtaking it in a sudden burst of speed, others chase prey over long distances and slowly tire it out. Some (such as skunks) simply shuffle about and eat whatever they happen to come across. Still others scavenge from carcasses or, in urban areas, from garbage cans. Some carnivores, such as arctic foxes (Vulpes lagopus) are known to cache their kills for later consumption. Crabeater seals (Lobodon carcinophaga) are unique among carnivores in that they are filter feeders. They have specialized teeth which allow them to strain tiny zooplankton, their staple food, from the water.
Foraging Behavior: stores or caches food ; filter-feeding
Primary Diet: carnivore (Eats terrestrial vertebrates, Piscivore , Eats eggs, Insectivore , Eats non-insect arthropods, Molluscivore , Scavenger ); herbivore (Folivore , Frugivore , Granivore ); omnivore ; planktivore
Carnivorans are important predators in many ecosystems, acting as a "top-down" control on populations of their prey. Many are such an important control on their prey that they act as keystone species, and their removal has drastic consequences for the ecosystem. For example, wolves were recently reintroduced to Yellowstone National Park after being extirpated for nearly 70 years, and their predation on elk has allowed woody plants to recover from overbrowsing.
Carnivores host a wide range of internal and external parasites, including protozoans, nematodes, trematodes, cestodes, fleas, lice, and ticks.
Ecosystem Impact: keystone species
- Ripple, W., R. Beschta. 2003. Wolf reintroduction, predation risk, and cottonwood recovery in Yellowstone National Park. Forest Ecology and Management, 184: 299-313.
- Roberts, L., J. Janovy Jr.. 2000. Foundations of Parasitology. New York: McGraw-Hill.
Many carnivorans are top predators in their ecosystems, and therefore do not face the threat of predation as adults, though their young may be vulnerable. Small terrestrial carnivorans face predation by larger carnivorans, and by diurnal and nocturnal birds of prey. Pinnipeds face predation by large cetaceans such as killer whales (Orcinus orca) and by sharks. Many carnivorans, large and small, terrestrial and aquatic, are hunted by humans.
Most carnivorans use their teeth and claws to fend off predators. A carnivore that feels threatened typically crouches and bares its teeth, hissing or growling at its attacker and biting and scratching if the attacker gets too close. Pinnipeds, on the other hand, rely largely on their speed and agility in the water to escape predators. Female carnivorans often hide their helpless infants in a den, and may switch the den location occasionally to avoid detection. Some carnivoran parents are also known to vigorously defend their offspring if necessary. Many carnivorans are the same color as their background (such as Arctic foxes, which turn white in winter to match the snow). They also frequently exhibit countershading or color patterns, such as spots and stripes, that break up their outline and make them difficult to see. A few carnivorans have special adaptations to defend themselves against predators. Skunks and some mustelids, herpestids, and viverrids have well-developed anal glands, which produce a foul-smelling musk that is released under stress. These animals usually bear aposematic coloration in the form of contrasting stripes and bands, warning would-be predators to stay away. Finally, it has been postulated that some carnivorans mimic others to avoid predation. For example, the coloration of cheetah cubs, which are highly vulnerable to predation, may mimic that of honey badgers, which are aposematic and highly aggressive.
- larger carnivores (Carnivora)
- diurnal birds of prey (Falconiformes)
- owls (Strigiformes)
- cetaceans (Cetacea)
- killer whales (Orcinus orca)
- sharks (Chondrichthyes)
- humans (Homo sapiens)
Anti-predator Adaptations: mimic; aposematic ; cryptic
- Eaton, R. 1976. A possible case of mimicry in larger mammals. Evolution, 30(4): 853-856.
- Korpimaki, E., K. Norrdahl. 1989. Avian predation on mustelids in Europe 1. Occurrence and effects on body size variation and life traits. Oikos, 55(2): 205-215.
- Ortolani, A., T. Caro. 1996. The adaptive significance of color patterns in carnivores: Phylogenetic tests of classic hypotheses. Pp. 132-188 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 2. Ithaca: Cornell University Press.
Known prey organisms
This list may not be complete but is based on published studies.
Life History and Behavior
Most carnivorans have acute senses. Vision and hearing are excellent in many, often far surpassing the capabilities of humans. Domestic cats and other carnivores that hunt small rodents can hear ultrasounds emitted by their prey. Most carnivorans come equipped with tactile hairs (vibrissae) on the face and legs, which they use to feel their way through narrow, dark surroundings. The sense of smell is often remarkable. Carnivorans make extensive use of chemical signals excreted in urine and feces and produced by glands in the skin and anal region. They use these chemical signals for scent-marking territories and for conveying information about identity, social status, and reproductive status. Carnivorans also communicate acoustically with a variety of yips, howls, barks, growls, roars, and purrs. These sounds have various functions, including strengthing social bonds, advertising for mates, defending territories, and communicating alarm, distress, and contentment. Social carnivores such as lions sometimes erupt into choruses of loud roars as a means of calling one another to assemble. Visual signals, mainly in the form of body posturing, are also used by carnivorans to communicate, and tactile signals, as when a wolf licks and bites the muzzle of a dominant individual to show submission, are used as well.
Communication Channels: visual ; tactile ; acoustic ; chemical
Other Communication Modes: choruses ; pheromones ; scent marks
Perception Channels: visual ; tactile ; acoustic ; ultrasound ; chemical
- Gorman, M., B. Trowbridge. 1989. The role of odor in the social lives of carnivores. Pp. 57-88 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
- Peters, G., W. Wozencraft. 1989. Acoustic communication by fissiped carnivores. Pp. 14-56 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
Carnivores are relatively long-lived mammals, with most species living at least a decade. The main exceptions are small weasels (Mustela), which live up to six years in captivity but usually do not make it past one year in the wild. In general, pinniped carnivores live longer than fissiped carnivores, with several species (walruses, gray seals, ringed seals, Caspian seals, and Baikal seals) reported to live 40 or more years in the wild.
- Carey, J., D. Judge. 2002. "Longevity Records: Life Spans of Mammals, Birds, Amphibians, Reptiles, and Fish" (On-line). Max Planck Institute for Demographic Research. Accessed December 05, 2005 at http://www.demogr.mpg.de/.
Carnivores have polygynous, polygynandrous, and monogamous mating systems. Southern elephant seals (Mirounga leonina) demonstrate extreme polygyny, wherein males fight for exclusive access to harems of females. Gray wolves (Canis lupus), on the other hand, are monogamous cooperative breeders; the dominant male and female of each pack breed and all members of the pack help raise their offspring. Solitary carnivores, such as bears, mustelids, and cats, are often polygynandrous, with males and females each having multiple partners during the breeding season.
Mating System: monogamous ; polygynous ; polygynandrous (promiscuous) ; cooperative breeder
Carnivores breed either aseasonally or seasonally; those in cold climates usually mate in winter and spring and give birth to their young during spring and summer. Females may be polyestrus or monoestrus; in some species, ovulation is induced by the act of mating. Carnivores may have two or three litters per year (as with least weasels), but most carnivore females have just one litter every one to two years. Delayed implantation, wherein the blastocyst lies quiescent for several months before implanting in the uterine lining, is common in some carnivore families (such as mustelids). After implantation, gestation periods range from five weeks in least weasels to 15 months in walruses. Typical true gestation periods last two to four months. Litter sizes range from 1 to 16, and are commonly 3 to 5. Females nurse their young for up to two years, and the young take up to seven years to reach sexual maturity.
Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; induced ovulation ; viviparous ; delayed implantation
Female carnivores nurture their young inside their bodies for up to 15 months and provide their young with milk after birth. The length of nursing varies considerably among carnivores. Some phocids only nurse their young for a couple of weeks, whereas walruses nurse their young for up to two years. The duration of lactation in terrestrial carnivores falls between these two extremes. Carnivore young range from highly precocial, harbor seal (Phoca vitulina) pups are able to swim a few minutes after birth, to altricial, as in bears. Female carnivores usually bear the sole responsiblity for nurturing and protecting their offspring, but male parental care is not uncommon, especially among canids. Carnivores that live in groups and breed communally may all share in the task of raising each others' offspring. In some social species, like spotted hyenas (Crocuta crocuta), the mother's position in the dominance hierarchy determines the position of her offspring. The young of spotted hyenas, wolverines, sea otters, bears, and large felids stay with their mothers for up to two years even though they are weaned well before this time; they depend on their mothers for food until they become proficient at hunting for themselves. In carnivores that form close-knit social groups, bonds between mother and offspring may extend well beyond the period of offspring dependency.
Parental Investment: altricial ; precocial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Male, Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning; inherits maternal/paternal territory; maternal position in the dominance hierarchy affects status of young
- Ewer, R. 1973. The carnivores. Ithaca: Cornell University Press.
- Frank, L. 1996. Female masculinization in the spotted hyena: Endocrinology, behavioral ecology, and evolution. Pp. 78-131 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 2. Ithaca: Cornell University Press.
- Gittleman, J. 1989. Carnivore group living: Comparative trends. Pp. 183-207 in J Gittleman, ed. Carnivore Behavior, Ecology and Evolution, vol. 1. Ithaca: Cornell University Press.
- King, C. 1989. The advantages and disadvantages of small size to weasels, Mustela species. Pp. 302-334 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
- Mead, R. 1989. The physiology and evolution of delayed implantation in carnivores. Pp. 437-464 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
- Moehlman, P. 1989. Intraspecific variation in canid social systems. Pp. 143-163 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
- Reeves, R., B. Stewart, P. Clapham, J. Powell. 2002. The National Audobon Society Guide to Marine Mammals of the World. New York: Alfred A. Knopf.
- Sandell, M. 1989. The mating tactics and spacing patterns of solitary carnivores. Pp. 164-182 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 1. Ithaca: Cornell University Press.
- Stains, H. 1984. Carnivores. Pp. 491-521 in S Anderson, J Jones Jr., eds. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons.
- Vaughan, T., J. Ryan, N. Czaplewski. 2000. Mammalogy, 4th Edition. Toronto: Brooks Cole.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:5854
Specimens with Barcodes:1967
Species With Barcodes:208
Members of Carnivora have been feared, persecuted, and exploited by humans for centuries. There are currently 122 species on the IUCN's Red List of Threatened Species. Of these, 11 are near threatened, 9 are lower risk, 39 are vulnerable, 33 are endangered, 6 are critically endangered, 5 have recently gone extinct, and one, black-footed ferrets (Mustela nigripes), is extinct in the wild, although reintroduction efforts show promise. Another 18 are listed as data deficient. Major threats to carnivorans include habitat loss and degradation and hunting for sport and profit. Rare species often fetch top dollar on the black market, even though trade in these species is strictly regulated by CITES and by national laws. Captive breeding programs may be the last chance for the survival of some species, such as giant pandas (Ailuropoda melanoleuca). In some cases, reintroduction of species into areas where they were previously extirpated has been successful, as with the wolves of Yellowstone. In order to save carnivorans from extinction in the long term, large swaths of habitat and healthy populations of prey species must be preserved in all parts of the world, and humans must learn to coexist peacefully with these animals.
- IUCN, 2004. "2004 IUCN Red List of Threatened Species" (On-line). Accessed December 06, 2005 at www.redlist.org.
Relevance to Humans and Ecosystems
Carnivorans also have negative impacts on humans. They may compete with humans for game and prey on livestock. Occasionally, large carnivorans even attack and kill humans. Omnivorous species may raid fruit crops, and in urban areas carnivorans become pests when they raid garbage cans and take up residence inside chimneys and under porches. They also carry diseases and parasites, such as rabies, that can be transmitted to humans and domestic animals.
Negative Impacts: injures humans (bites or stings, carries human disease); crop pest; causes or carries domestic animal disease ; household pest
Humans benefit from carnivorans in many ways. Humans have hunted them for thousands of years for sport and for their fur, meat, and other body parts. Bones and soft tissues of tigers (Panthera tigris) and other large carnivores have long been used in traditional Asian medicine. Millions of small carnivores such as red foxes (Vulpes vulpes) and mink (Neovison vison) are raised on farms for their fur. Carnivorans are also valuable to humans for their ability to control rodents and other pests, and domesticated cats, dogs, and other carnivorans are popular pets worldwide.
Positive Impacts: pet trade ; food ; body parts are source of valuable material; ecotourism ; source of medicine or drug ; research and education; controls pest population
- Schaller, G. 1996. Introduction: Carnivores and conservation biology. Pp. 1-10 in J Gittleman, ed. Carnivore Behavior, Ecology, and Evolution, vol. 2. Ithaca: Cornell University Press.
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