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Reproduction

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Behavior

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Conservation Status

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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.

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Comprehensive Description

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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.

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Benefits

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Benefits

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Associations

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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

Commensal/Parasitic Species:

  • protozoans
  • nematodes (Nematoda)
  • trematodes (Trematoda)
  • cestodes (Cestoda)
  • fleas (Siphonaptera)
  • lice (Anoplura)
  • ticks (Acari)
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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Trophic Strategy

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Distribution

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Habitat

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Life Expectancy

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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.

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Morphology

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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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Allison Poor, University of Michigan-Ann Arbor
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Associations

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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.

Known Predators:

  • 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

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Poor, A. . "Carnivora" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Carnivora.html
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Brief Summary

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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.

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Carnivora

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Carnivora (/kɑːrˈnɪvərə/;[3][4] ; from Latin carō (stem carn-) "flesh" and vorāre "to devour") is a clade of placental mammals where most species have specialized in primarily eating flesh. However some species are omnivorous, like raccoons and bears, and quite few species like pandas are specialized herbivores. Its members are formally referred to as carnivorans, whereas the word "carnivore" (often popularly applied to members of this group) can refer to any meat-eating organism. With at least 279 extant species found on every major landmass and in a variety of habitats, ranging the cold polar regions to the hyper-arid region of the Sahara Desert to the open seas even, Carnivora is the fifth largest order of mammals and one of the more successful members of the group. They come in a huge array of different body plans in contrasting shapes and sizes. The smallest carnivoran is the least weasel (Mustela nivalis), which can weigh as little as 25 g (0.88 oz) and 11 cm (4.3 in), as the largest is the southern elephant seal (Mirounga leonina), whose adult males weigh up to 5,000 kg (11,000 lb) and measure up to 6.7 m (22 ft) in length. All species of carnivorans are descended from a group of mammals which were related to today's pangolins, having appeared in North America 6 million years after the Cretaceous–Paleogene extinction event.[5][6] These early ancestors of carnivorans would have resembled small weasel or genet-like mammals, occupying a nocturnal shift on the forest floor or in the trees, as other groups of mammals like the mesonychians and creodonts were occupying the top faunivorous niche. However, by the time Miocene epoch appeared, most if not all of the major lineages and families of carnivorans had diversified and took over this niche.

Carnivora can be divided into two subclades: the cat-like Feliformia and the dog-like Caniformia which are differentiated based on the structure of their ear bones and cranial features. The feliforms include families such as the cats, the hyenas, the mongooses and the civets. Majority of feliform species are found in the Old World, though the cats and one extinct genus of hyena have successfully diversified into the Americas. The caniforms include the dogs, bears, raccoons, weasels, and pinnipeds. Members of this group are found worldwide and with incredible diversity in their diet, behavior, and morphology. Despite this the two groups of carnivorans share several unique traits, one being the presence of the carnassial teeth. In carnivorans the carnassial pair is made up by the fourth upper premolar and the first lower molar teeth. There is variation among the carnassial pair depending on the family. Some species are cursorial and the foot posture in terrestrial species is either digitigrade or plantigrade. In pinnipeds the feet have become flippers where the locomotion is unique in each of the pinniped families.

Carnivorans are arguably after primates the group of mammals most fascinated or of interest to humans. The dog is noteworthy for not only being the first species of carnivoran to be domesticated, but also the first species of any organism. In the last 10 to 12,000 years humans have selectively bred dogs for a variety of different tasks and today there are well over 400 breeds. The cat is another domesticated carnivoran and it is today considered one of the most successful species on the planet, due to their close proximity to humans and the popularity of cats as pets. Many other species popular, often part of the charismatic megafauna where every major civilization has incorporated a species of carnivoran into their culture such as the lion, viewed as royalty. Yet many species such as wolves and the big cats have been hunted and persecution, with some places they have been extirpated. Habitat loss and human encroachment as well as climate change have been the primary cause of many species to go into decline. Most species need large adequate amount of territory and plenty of food resources in order to survive, and the decline of both of them are the primary reasons of decline. Four species of carnivorans have gone extinct since the 1600s: Falkland Island Wolf (Dusicyon australis) in 1876; the Sea Mink (Neovison macrodon) in 1894; the Japanese Sea Lion (Zalophus japonicus) in 1951 and the Caribbean Monk Seal (Neomonachus tropicalis) in 1952.[5] Some species such as the red fox (Vulpes vulpes) and stoat (Mustela erminea) have been introduced to Australasia that have caused many native species to become endangered or even extinct.[7]

Systematics

Evolution

"
Life reconstruction of Tapocyon robustus, a species of miacid

The order Carnivora belongs to a group of mammals known as Laurasiatheria, which also includes other groups such as bats and ungulates.[8][9] Within this group the carnivorans are placed in the clade Ferae. Ferae includes the closest extant relative of carnivorans, the pangolins, as well as several extinct groups of mostly Paleogene carnivorous placentals such as the creodonts, the arctocyonians, and mesonychians.[10] The creodonts were originally thought of as the sister taxon to the carnivorans, perhaps even ancestral to, based on the presence of the carnassial teeth.[11] but the nature of the carnassial teeth is different between the two groups. In carnivorans the carnassials are positioned near the front of the molar row, while in the creodonts they are positioned near the back of the molar row.[12] and this suggests a separate evolutionary history and an order-level distinction.[13] In addition recent phylogenetic analysis suggests that creodonts are more closely related to pangolins while mesonychians might be the sister group to carnivorans and their stem-relatives.[10]

The closest stem-carnivorans are the miacoids. The miacoids include the families Viverravidae and Miacidae, and together the Carnivora and Miacoidea form the stem-clade Carnivoramorpha. The miacoids were small, gennet-like carnivoramorphs that occupy a variety of niches such as terrestrial and arboreal habitats. Recent studies have shown a supporting amount of evidence that Miacoidea is an evolutionary grade of canivoramorphs that, while viverravids are monophyletic basal group, the miacids are paraphyletic in respect to Carnivora (as shown in the phylogeny below).[14][15][16][17][18][19]

Ferae

Pholidotamorpha "FMIB

  †Oxyaenodonta

Oxyaenidae "Oxyaena

   

Hyaenodonta "Hyaenodon

    Carnivoraformes    

Neovulpavus

   

"Miacis" medius

               

Dormaalocyon

   

"Miacis" exiguus

       

Vassacyon

   

Vulpavus

       

"Miacis" deutschi

    †Uintacyon

Miocyon

?

Simamphicyon

   

Zodiocyon

   

Uintacyon (sensu stricto)

sensu lato

Dawsonicyon

   

Miacis

   

miacid sp. (PM 3868)

?

Eosictis

   

"Miacis" petilus

   

"Miacis" latidens

?

Eogale

?

Chailicyon

    ?

Paroodectes

?

Paramiacis

     

Gracilocyon

   

Oodectes

         

Ceruttia

     

Prodaphaenus

       

Procynodictis

   

Harpalodon

     

Walshius

     

Tapocyon

    †Quercygalidae

Quercygale

?

Messelogale

    Carnivora

Feliformia "Stamp-russia2014-save-russian-cats-(snow

   

Caniformia "Dogs,

sensu stricto          
(Carnivora sensu lato)

Carnivoramorpha as a whole first appeared in the Paleocene of North America about 60 million years ago.[6] Crowned carnivorans first appeared around 42 million years ago in the Middle Eocene.[1] Their molecular phylogeny shows the extant Carnivora are a monophyletic group, the crown group of the Carnivoramorpha.[20] From there carnivorans have split into two clades based on the composition of the bony structures that surround the middle ear of the skull, the cat-like feliforms and the dog-like caniforms.[21] In feliforms, the auditory bullae are double-chambered, composed of two bones joined by a septum. Caniforms have single-chambered or partially divided auditory bullae, composed of a single bone.[22] Initially the early represents of carnivorans were small as the creodonts dominated the niches as top apex predators, but by the Miocene most of the extant carnivoran families have diversified and successfully out-competed the creodonts.

The phylogenetic relationships of the carnivorans are shown in the following cladogram:[23][24][25][26][27]

Carnivoraformes

Miacidae (paraphyletic family) "Miacis

   

Ceruttia

   

Walshius

Carnivora FeliformiaAeluroideaViverroidea Herpestoidea Hyaenidae

Percrocutidae "Dinocrocuta

   

Hyaenidae (hyaenas) "Hyaena

   

Lophocyonidae

sensu lato Herpestidae

Herpestidae (mongoose) "Lydekker

   

Eupleridae (Malagasy mongooses) "Cryptoprocta

sensu lato    

Viverridae (viverrids) "Malay

         

Shandgolictis

   

Asiavorator

   

Alagtsavbaatar

     

Anictis

FeloideaPrionodontidae

Prionodontidae (Asiatic linsangs) "Prionodon

   

Palaeoprionodon

sensu lato

Haplogale

     

Stenoplesictis

?

Pseudictis

Felidae

Barbourofelidae

     

Viretictis

     

Stenogale

   

Felidae (cats) "Stamp-russia2014-save-russian-cats-(snow

      sensu lato        

Nandiniidae (African palm civet) "The

     

Nimravidae (false saber-toothed cats) "Dinictis

     

Palaeogalidae

    Caniformia   †Amphicyonoidea

Amphicyonidae ("bear-dogs") "Ysengrinia.jpg"

       

Lycophocyon

Canoidea Cynoidea

Canidae (canids) "Dogs,

ArctoideaUrsida Ursoidea

Ursidae (bears) "Ursus

   

Adracon

        Musteloidea        

Procyonidae (raccoons) "Wild

   

Mustelidae (mustelids) "Fitch

     

Ailuridae "RedPandaFullBody

    ?

Peignictis

   

Mephitidae (skunks) "Die

     

Plesiogale

    Pinnipedimorpha

Semantoridae

     

Enaliarctidae

     

Pinnarctidion

Pinnipediformes    

Pacificotaria

   

Pteronarctos

    Pinnipedia Otarioidea

Otariidae
(eared seals)

"Zalophus

   

Odobenidae
(walruses)

"USSR

    Phocoidea

Desmatophocidae

   

Phocidae
(earless seals)

"Faroe

    sensu stricto      
(Pinnipedia sensu lato)
             

Classification of the extant carnivorans

In 1758 the Swedish botanist Carl Linnaeus placed all the carnivorans that was known at the time in the group Ferae (not to be confused with the modern concept of Ferae which also include pangolins) in the tenth edition of his book Systema Naturae. He recongized six genera: Canis (canids and hyaenids), Phoca (pinnipeds), Felis (felids), Viverra (viverrids, herpestids, and mephitids), Mustela (non-badger mustelids), Ursus (ursids, large species of mustelids, and procyonids).[28] It wasn't until in 1821 when the English writer and traveler Thomas Edward Bowdich gave the group its modern and accepted name.[2]

Initially the modern concept of Carnivora was divided into two suborders: the terrestrial Fissipedia and the marine Pinnipedia.[29] Below is the classification of how the extant families were related to each other after American paleontologist George Gaylord Simpson in 1945:[29]

  • Order Carnivora Bowdich, 1821
    • Suborder Fissipedia Blumenbach, 1791
      • Superfamily Canoidea G. Fischer de Waldheim, 1817
        • Family Canidae G. Fischer de Waldheim, 1817 – dogs
        • Family Ursidae G. Fischer de Waldheim, 1817 – bears
        • Family Procyonidae Bonaparte, 1850 – raccoons and pandas
        • Family Mustelidae G. Fischer de Waldheim, 1817 – skunks, badgers, otters and weasels
      • Superfamily Feloidea G. Fischer de Waldheim, 1817
        • Family Viverridae J. E. Gray, 1821 – civets and mongooses
        • Family Hyaenidae J. E. Gray, 1821 – hyenas
        • Family Felidae G. Fischer de Waldheim, 1817 – cats
    • Suborder Pinnipedia Iliger, 1811
      • Family Otariidae J. E. Gray, 1825 – eared seals
      • Family Odobenidae J. A. Allen, 1880 – walrus
      • Family Phocidae J. E. Gray, 1821 – earless seals

Since then, however, the methods in which mammalogists use to assess the phylogenetic relationships among the carnivoran families has been improved with using more complicated and intensive incorporation of genetics, morphology and the fossil record. Research into Carnivora phylogeny since 1945 has found Fisspedia to be paraphlyetic in respect to Pinnipedia,[30] with pinnipeds being either more closely related to bears or to weasels.[31][32][33][34][35] The small carnivoran families Viverridae,[36] Procyonidae, and Mustelidae have found to be polyphyletic:

  • Mongooses and a handful of Malagasy endemic species are found to be in a clade with hyenas, which the Malagasy species being in their own family Eupleridae.[37][38][39]
  • The African palm civet is a basal cat-like carnivoran.[40]
  • The linsang is more closely related to cats.[41]
  • Pandas are not procyonids nor are they are natural grouping.[42] The giant panda is a true bear[43][44] while the red panda is a distinct family.[45]
  • Skunks and stink badgers are placed in their own family, and are the sister group to a clade pertaining Ailuridae, Procyonidae and Mustelidae sensu stricto.[46][45]

Below is a table chart of the extant carnivoran families and number of extant species recognized by various authors of the first and fourth volumes of Handbook of the Mammals of the World published in 2009[47] and 2014[48] respectively:

Anatomy and physiology

Craniodental region

"
A skull of a fossa (Cryptoprocta ferox), note the large and conical canine and carnassial teeth common in feliforms.

The canine teeth are usually large and conical. The canines are thick and incredibly stress resistant. All of the terrestrial species of carnivorans have three incisors on the top and bottom row of the dentition (the exception being is the sea otter (Enhydra lutris) which only has two lower incisor teeth).[49][50] The third molar has been lost. The carnassial pair is made up by the fourth upper premolar and the first lower molar teeth. Like most mammals the dentition is heterodont in nature, though in some species like the aardwolf (Proteles cristata) the teeth have been greatly reduced and the cheek teeth are specialised for eating insects. In pinnipeds the teeth are homodont as they have evolved to grasp or to catch fish, and the cheek teeth are often lost.[50] In bears and raccoons the carnassial pair is secondarily reduced.[50] The skulls are heavily built with a strong zygomatic arch.[50] Often a sagittal crest is present, sometimes more evident in sexual dimorphic species like sea lions and fur seals, though it has also been greatly reduced seen in some small carnivorans.[50] The braincase is enlarged and the frontoparietal is position at the front of it. In most species the eyes are position at the front of the face. In caniforms the rostrum is usually longer with many teeth, where in comparison with felifoms the rostrum is shorter and have fewer teeth. The carnassial teeth in feliforms, however is more sectional.[50] The tubinares are large and complex in comparison to other mammals, providing a large surface area for olfactory receptors.[50]

Postcranial region

"
A black-backed jackal (Canis mesomelas) trying to predate on a brown fur seal (Arctocephalus pusillus) pup. These two species illustrate the diversity in bodyplan seen among carnivorans, especially between pinnipeds and their terrestrial relatives.

Aside from an accumulation of characteristics in the dental and cranial features, not much of their overall anatomy unites them as a group.[49] All species of carnivorans have quadrupedal limbs with usually five digits at the front feet and four digits at the back feet. In terrestrial carnivorans the feet have soft pads. The feet can either be digitgrade seen in cats, hyenas and dogs or plantigrade seen in bears, skunks, raccoons, weasels, civets and mongooses. In pinnipeds the limbs have been modified into flippers. Unlike other marine mammals, such as cetaceans and sirenians which have fully functional tails to help them swim, pinnipeds use their limbs underwater for locomotion. In earless seals they use their back flippers; sea lions and fur seals use their front flippers, and the walrus use all of their limbs. This resulted in pinnipeds having significantly shorter tails. Aside from the pinnipeds, dogs, bears, hyenas, and cats have distinct and recognizable appearances. Dogs are usually cursorial mammals and are gracile in appearance, often relying on their teeth to hold to prey; bears are much larger and rely on their physical strength to forage for food. Cats in comparison to dogs and bears have much longer and stronger frontlimbs armed with retractable claws to hold on to prey. Hyenas are dog-like feliforms that have sloping backs due to their front legs being longer than their hindlegs. The raccoon family as well as the red panda are small, bear-like carnivorans with long tails. The other small carnivoran families Nandiniidae, Prionodontidae, Viverridae, Herpestidae, Eupleridae, Mephitidae and Mustelidae have through convergent evolution maintained the small, ancestral appearance of the miacoids, though there is some variation seen such as the robust and stout physicality of badgers and the wolverine (Gulo gulo).[49]

Depending on the environment the species is, the length and density of their fur varies. In warm climate species the fur is often short in length and lighter. In comparison to cold climate species the fur is either dense or long, often with an oily substance to keep them warm. The pelage coloration comes in many colors, often including black, white, orange, yellow, red, and many shades of gray and brown. There can be colored patterns too, such striped, spotted, blotched, banded, or otherwise boldly patterned. There seems to be a correlation between habitat and color pattern as for example spotted or banded species tend to be found in heavily forested environments.[49] Some species like the grey wolf is a polymorphic species with different individual variation in colors. The arctic fox (Vulpes lagopus) and the stoat (Mustela erminea) the fur goes from white and dense in the winter to brown and sparse in the summer. In pinnipeds, polar bears, and sea otters have a thick insulating layer of blubber to help maintain their body temperature.

See also

References

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Carnivora: Brief Summary

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Carnivora (/kɑːrˈnɪvərə/; ; from Latin carō (stem carn-) "flesh" and vorāre "to devour") is a clade of placental mammals where most species have specialized in primarily eating flesh. However some species are omnivorous, like raccoons and bears, and quite few species like pandas are specialized herbivores. Its members are formally referred to as carnivorans, whereas the word "carnivore" (often popularly applied to members of this group) can refer to any meat-eating organism. With at least 279 extant species found on every major landmass and in a variety of habitats, ranging the cold polar regions to the hyper-arid region of the Sahara Desert to the open seas even, Carnivora is the fifth largest order of mammals and one of the more successful members of the group. They come in a huge array of different body plans in contrasting shapes and sizes. The smallest carnivoran is the least weasel (Mustela nivalis), which can weigh as little as 25 g (0.88 oz) and 11 cm (4.3 in), as the largest is the southern elephant seal (Mirounga leonina), whose adult males weigh up to 5,000 kg (11,000 lb) and measure up to 6.7 m (22 ft) in length. All species of carnivorans are descended from a group of mammals which were related to today's pangolins, having appeared in North America 6 million years after the Cretaceous–Paleogene extinction event. These early ancestors of carnivorans would have resembled small weasel or genet-like mammals, occupying a nocturnal shift on the forest floor or in the trees, as other groups of mammals like the mesonychians and creodonts were occupying the top faunivorous niche. However, by the time Miocene epoch appeared, most if not all of the major lineages and families of carnivorans had diversified and took over this niche.

Carnivora can be divided into two subclades: the cat-like Feliformia and the dog-like Caniformia which are differentiated based on the structure of their ear bones and cranial features. The feliforms include families such as the cats, the hyenas, the mongooses and the civets. Majority of feliform species are found in the Old World, though the cats and one extinct genus of hyena have successfully diversified into the Americas. The caniforms include the dogs, bears, raccoons, weasels, and pinnipeds. Members of this group are found worldwide and with incredible diversity in their diet, behavior, and morphology. Despite this the two groups of carnivorans share several unique traits, one being the presence of the carnassial teeth. In carnivorans the carnassial pair is made up by the fourth upper premolar and the first lower molar teeth. There is variation among the carnassial pair depending on the family. Some species are cursorial and the foot posture in terrestrial species is either digitigrade or plantigrade. In pinnipeds the feet have become flippers where the locomotion is unique in each of the pinniped families.

Carnivorans are arguably after primates the group of mammals most fascinated or of interest to humans. The dog is noteworthy for not only being the first species of carnivoran to be domesticated, but also the first species of any organism. In the last 10 to 12,000 years humans have selectively bred dogs for a variety of different tasks and today there are well over 400 breeds. The cat is another domesticated carnivoran and it is today considered one of the most successful species on the planet, due to their close proximity to humans and the popularity of cats as pets. Many other species popular, often part of the charismatic megafauna where every major civilization has incorporated a species of carnivoran into their culture such as the lion, viewed as royalty. Yet many species such as wolves and the big cats have been hunted and persecution, with some places they have been extirpated. Habitat loss and human encroachment as well as climate change have been the primary cause of many species to go into decline. Most species need large adequate amount of territory and plenty of food resources in order to survive, and the decline of both of them are the primary reasons of decline. Four species of carnivorans have gone extinct since the 1600s: Falkland Island Wolf (Dusicyon australis) in 1876; the Sea Mink (Neovison macrodon) in 1894; the Japanese Sea Lion (Zalophus japonicus) in 1951 and the Caribbean Monk Seal (Neomonachus tropicalis) in 1952. Some species such as the red fox (Vulpes vulpes) and stoat (Mustela erminea) have been introduced to Australasia that have caused many native species to become endangered or even extinct.

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