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Temporal range: 42-0 Ma
Middle Eocene-Holocene[1]
Carnivora portraits.jpg
Various carnivorans, with feliforms to the left, and caniforms to the right
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
(unranked): Carnivoraformes
Order: Carnivora
Bowdich (1821)[2]

Carnivora (;[3][4] ; from Latin car? (stem carn-) "flesh" and vor?re "to devour") is a diverse scrotiferan order that includes over 280 species of placental mammals. 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. Carnivorans are the most diverse in size of any mammalian order, ranging from the least weasel (Mustela nivalis), at as little as 25 g (0.88 oz) and 11 cm (4.3 in), to the polar bear (Ursus maritimus), which can weigh up to 1,000 kg (2,200 lb), to 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.

Carnivorans have teeth and claws adapted for catching and eating other animals. Many hunt in packs and are social animals, giving them an advantage over larger prey. Some carnivorans, such as cats and pinnipeds, depend entirely on meat for their nutrition. Others, such as raccoons and bears, are more omnivorous, depending on the habitat. The giant panda is largely a herbivore, but also feeds on fish, eggs, and insects. The polar bear subsists mainly on seals.

Carnivorans are split into two suborders: Feliformia ("catlike") and Caniformia ("doglike").


Carnivore (wolf) mandible diagram showing the names and positions of the teeth

Carnivorans all share the same arrangement of teeth in which the last upper premolar (named P4) and the first lower molar (named m1) have blade-like enamel crowns that work together as carnassial teeth to shear meat. Carnivorans have had this arrangement for over 60 million years with many adaptions, and these dental adaptions help identify carnivoran species and groupings of species.[5]


Carnivorans evolved in North America out of members of the family Miacidae (miacids) about 42 million years ago.[1] The split into cat-like and dog-like forms (Feliformia and Caniformia) occurred relatively soon after. Their molecular phylogeny[6] shows the extant Carnivora are a monophyletic group, the crown group of the Carnivoramorpha.

Distinguishing features


Most carnivorans are terrestrial; they usually have strong, sharp claws, typically with five, but never fewer than four, toes on each foot, and well-developed, prominent canine teeth, cheek teeth (premolars, and molars) that generally have cutting edges.[7] The last premolar of the upper jaw and first molar of the lower are termed the carnassials or sectorial teeth. These blade-like teeth occlude (close) with a scissor-like action for shearing and shredding meat. Carnassials are most highly developed in the Felidae and the least developed in the Ursidae. Carnivorans have six incisors and two conical canines in each jaw. The only two exceptions to this are the sea otter (Enhydra lutris), which has four incisors in the lower jaw, and the sloth bear (Melursus ursinus), which has four incisors in the upper jaw. The number of molars and premolars is variable between carnivoran species, but all teeth are deeply rooted and are diphyodont.[8] Incisors are retained by carnivorans and the third incisor is commonly large and sharp (canine-like). Carnivorans have either four or five digits on each foot, with the first digit on the forepaws, also known as the dew claw, being vestigial in most species and absent in some.[]

The superfamily Canoidea (or suborder Caniformia) - Canidae (wolves, dogs and foxes), Mephitidae (skunks and stink badgers), Mustelidae (weasels, badgers, and otters), Procyonidae (raccoons), Ursidae (bears), Ailuridae (red panda), Otariidae (eared seals), Odobenidae (walrus), and Phocidae (earless seals) (the last three families formerly classified in the superfamily Pinnipedia) and the extinct family Amphicyonidae (bear-dogs) - are characterized by having nonchambered or partially chambered auditory bullae, nonretractable claws, and a well-developed baculum. Most species are rather plain in coloration, lacking the flashy spotted or rosetted coats like many species of felids and viverrids have. This is because Canoidea tend to range in the temperate and subarctic biomes, although Mustelidae and Procyonidae have a few tropical species. Most are terrestrial, although a few species, like procyonids, are arboreal. All families except the Canidae and a few species of Mustelidae are plantigrade. Diet is varied and most tend to be omnivorous to some degree, and thus the carnassial teeth are less specialized. Canoidea have more premolars and molars in an elongated skull.

The superfamily Feloidea (or suborder Feliformia)- Felidae (cats), Prionodontidae (Asiatic linsangs), Herpestidae (mongooses), Hyaenidae (hyenas), Viverridae (civets), and Eupleridae (Malagasy carnivorans), as well as the extinct family Nimravidae (paleofelids) - often have spotted, rosetted or striped coats, and tend to be more brilliantly colored than their Canoidean counterparts. This is because these species tend to range in tropical habitats, although a few species do inhabit temperate and subarctic habitats. Many are arboreal or semiarboreal, and the majority are digitigrade. Diet tends to be more strictly carnivorous, especially in the family Felidae. They have fewer teeth and shorter skulls, with much more specialized carnassials meant for shearing meat. Feliformia claws are often retractile, or rarely, semiretractile. The terminal phalanx, with the claw attached, folds back in the forefoot into a sheath by the outer side of the middle phalanx of the digit, and is retained in this position when at rest by a strong elastic ligament. In the hindfoot, the terminal joint or phalanx is retracted on to the top, and not the side of the middle phalanx. Deep flexor muscles straighten the terminal phalanges, so the claws protrude from their sheaths, and the soft "velvety" paw becomes suddenly converted into a formidable weapon. The habitual retraction of the claws preserves their points from wear.[]


Skull structure

Skull of Hyaena eximia
Diagram of a wolf skull with key features labelled




Dentition relates to the arrangement of teeth in the mouth, with the dental notation for the upper-jaw teeth using the upper-case letters I to denote incisors, C for canines, P for premolars, and M for molars, and the lower-case letters i, c, p and m to denote the mandible teeth. Teeth are numbered using one side of the mouth and from the front of the mouth to the back. In carnivores, the upper premolar P4 and the lower molar m1 form the carnassials that are used together in a scissor-like action to shear the muscle and tendon of prey.[9]



Diet specializations


Reproductive system

A cat with a litter of four kittens

Most male carnivorans have a baculum, although it is relatively short in felids, and absent in hyenas.[10] Carnivorans tend to produce a single litter annually,[8] but some produce multiple litters a year, and larger carnivorans, like bears, have gaps of 2-3 yr between litters. The average gestation period lies between 50 and 115 days, although the ursids and mustelids have delayed implantation, thus extending the gestation period six to nine months beyond the normal period. Litter sizes are usually small, ranging from one to 13 young, which are born with underdeveloped eyes and ears. In most species, the mother has exclusive or at least primary care of the offspring. Many species of carnivorans are solitary, but a few are gregarious.[]


Artist's impression of one of the first carnivorans, a miacid

Carnivorans evolved from members of the family Miacidae (miacids, now recognized as paraphyletic[11]). The transition from Miacidae to Carnivora was a general trend in the middle and late Eocene, with taxa from both North America and Eurasia involved. The divergence of carnivorans from other miacids, as well as the divergence of the two clades within Carnivora, Caniformia and Feliformia, is now inferred to have happened in the middle Eocene, about 42 million years ago (mya).[1]




An African palm civet, possibly an example of a primitive feliform

The ancestor of all Feliformia evolved from the Caniformia-Feliformia split. Nandinia, the African palm civet, seems to be the most primitive of all the feliforms and the first to split from the others.[6] The Asiatic linsangs of the genus Prionodon (traditionally placed in the Viverridae) form their own family, as some recent[16] studies indicate that Prionodon is actually the closest living relative to the cats. The Nimravidae are sometimes seen as the most basal of all feliforms and the first to split from the others, but there is a possibility that Nimravidae might not even belong within the order,[15] and therefore its position as a clade within Carnivora is currently unstable. Other studies indicate that the barbourofelids form a separate family, which is closely related to the true felids instead of being related to the nimravids. Recognizable nimravid fossils date from the late Eocene (37 mya), from the Chadronian White River Carnivora Formation at Flagstaff Rim, Wyoming. Nimravid diversity appears to have peaked about 28 mya. The hypercarnivorous (strictly meat-eating) nimravid feliforms were extinct in North America after 26 mya and felids did not arrive in North America until the early middle Miocene (16 mya).


A recent study has finally resolved the exact position of Ailurus: the red panda is neither a procyonid nor an ursid, but forms a monotypic family, with the other musteloids as its closest living relatives. The same study also showed that the mustelids are not a primitive family, as was once thought. Their small body size is a secondary trait--the primitive body form of the arctoids was large, not small.[14] Recent molecular studies also suggest that the endemic Carnivora of Madagascar, including three genera usually classed with the civets and four genera of mongooses classed with the Herpestidae, are all descended from a single ancestor. They form a single sister taxon to the Herpestidae. The hyenas are also closely related to this clade.


Least weasel, the smallest carnivoran
Southern elephant seal, the largest carnivoran
Brown bear, one of the largest terrestrial carnivorans
A cat and a dog, domesticated carnivorans

When the order Carnivora was first named, there were only five families:


Phylogenetic tree

The phylogenetic relationships of carnivora are shown in the following cladogram:[18][19][21][22][23]




+Percrocutidae Dinocrocuta gigantea.jpg

Herpestidae (mongoose) Lydekker - Broad-banded Cusimanse (white background).JPG

Eupleridae (Malagasy mongooses) Cryptoprocta ferox - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).png

Viverridae (viverrids) Viverra civetta - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg


Prionodontidae (Asiatic linsang) Spotted linsang


Felidae (cats) Wildcat

Nandiniidae (African palm civet) Two-spotted palm civet

+Nimravidae (false saber-toothed cats) Dinictis


+Amphicyonidae ("bear-dogs") Ysengrinia americana



Canidae (canids) Dogs, jackals, wolves, and foxes (Plate XI).jpg


+Hemicyonidae ("dog-bears") Hemicyon white background.jpg

Ursidae (bears) Ursus thibetanus - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg


Otariidae (eared seals) Zalophus californianus J. Smit (white background).jpg

Odobenidae (walruses) USSR stamp Walrus 1977 (white background).png

Phocidae (earless seals) Faroe stamp 227 grey seal (Phoca vitulina) white background.jpg



Ailuridae Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg

Mephitidae (skunks) Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate CXXI-) (white background).jpg

Procyonidae (raccoons) Wild animals of North America, intimate studies of big and little creatures of the mammal kingdom (Page 410) (white background).jpg

Mustelidae (mustelids) European polecat

See also


  1. ^ a b c Heinrich, R.E.; Strait, S.G.; Houde, P. (January 2008). "Earliest Eocene Miacidae (Mammalia: Carnivora) from northwestern Wyoming". Journal of Paleontology. 82 (1): 154-162. doi:10.1666/05-118.1.
  2. ^ Bowditch, T. E. 1821. An analysis of the natural classifications of Mammalia for the use of students and travelers J. Smith Paris. 115. (refer pages 24, 33)
  3. ^ "Carnivora". Dictionary.com Unabridged. Random House.
  4. ^ "Carnivora". Merriam-Webster Dictionary.
  5. ^ Wang, Xiaoming; Tedford, Richard H.; Dogs: Their Fossil Relatives and Evolutionary History. New York: Columbia University Press, 2008. ISBN 0231135289, p. 1
  6. ^ a b Eizirik, E.; Murphy, W.J.; Koepfli, K.P.; Johnson, W.E.; Dragoo, J.W.; O'Brien, S.J. (July 2010). "Pattern and timing of the diversification of the mammalian order Carnivora inferred from multiple nuclear gene sequences". Molecular Phylogenetics and Evolution. 56 (1): 49-63. doi:10.1016/j.ympev.2010.01.033. PMID 20138220.
  7. ^ "The Teeth of Herbivores, Carnivores and Omnivores". Miami Center for Cosmetic and Implant Dentistry. Retrieved .
  8. ^ a b c Animal Diversity[better source needed]
  9. ^ Wang, Xiaoming; Tedford, Richard H. (2008). Dogs: Their Fossil Relatives and Evolutionary History. Columbia University Press, New York. p. 74. ISBN 978-0-231-13529-0.
  10. ^ Dixson, A. F. "Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)." Journal of Zoology 235.1 (1995): pp. 67-76. Archived June 3, 2016, at the Wayback Machine
  11. ^ Wesley-Hunt, G.D.; Flynn J.J. (2005). Phylogeny of the Carnivora: Basal Relationships Among the Carnivoramorphans, and Assessment of the Position of 'Miacoidea' Relative to Carnivora. Journal of Systematic Palaeontology, 3: 1-28. doi:10.1017/S1477201904001518
  12. ^ Polly, David; Gina D. Wesley-Hunt; Ronald E. Heinrich; Graham Davis & Peter Houde (September 2006). "Earliest Known Carnivoran Auditory Bulla and Support for a Recent Origin of Crown-Clade Carnivora (Eutheria, Mammalia)" (PDF). Palaeontology. 49 (5): 1019-1027. doi:10.1111/j.1475-4983.2006.00586.x. Archived (PDF) from the original on 24 May 2011.
  14. ^ a b Wesley-Hunt, Gina D.; John J. Flynn (2005). "Phylogeny of the Carnivores". Journal of Systematic Palaeontology. 3: 1-28. doi:10.1017/S1477201904001518. Archived from the original on 19 November 2010.
  15. ^ a b Wesley-Hunt, Gina D.; Lars Werdelin (2005). "Basicranial morphology and phylogenetic position of the upper Eocene carnivoramorphan Quercygale" (PDF). Acta Palaeontologica Polonica. 50 (4): 837-846. Archived from the original (PDF) on September 27, 2007.
  16. ^ Johnson, W.E.; Eizirik, E.; Pecon-Slattery, J.; Murphy, W.J.; Antunes, A.; Teeling, E.; O'Brien, S.J. (6 January 2006). "The Late Miocene Radiation of Modern Felidae: A Genetic Assessment". Science. 311 (5757): 73-77. Bibcode:2006Sci...311...73J. doi:10.1126/science.1122277. PMID 16400146.
  17. ^ Wesley-Hunt, Gina D. (2005). "The Morphological Diversification of Carnivores in North America". Paleobiology. 31: 35-55. doi:10.1666/0094-8373(2005)031<0035:TMDOCI>2.0.CO;2. ISSN 0094-8373. Archived from the original on 2008-12-05.
  18. ^ a b Wilson, D.E.; Mittermeier, R.A., eds. (2009). Handbook of the Mammals of the World, Volume 1: Carnivora. Barcelona: Lynx Ediciones. pp. 50-658. ISBN 978-84-96553-49-1.
  19. ^ a b c Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D. W.; Loveridge, A. J. (eds.). Biology and Conservation of Wild Felids. Oxford, UK: Oxford University Press. pp. 59-82. ISBN 978-0-19-923445-5.
  20. ^ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 548-559. ISBN 978-0-8018-8221-0. OCLC 62265494.
  21. ^ Flynn, J. J.; Finarelli, J. A.; Zehr, S.; Hsu, J.; Nedbal, M. A. (April 2005). "Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships". Systematic Biology. 54 (2): 317-37. doi:10.1080/10635150590923326. PMID 16012099.
  22. ^ Morales, Jorge; Mayda, Serdar; Valenciano, Alberto; DeMiguel, Daniel; Kaya, Tanju (2019). "A new lophocyonid, Izmirictis cani gen. et sp. nov. (Carnivora: Mammalia), from the lower Miocene of Turkey". Journal of Systematic Palaeontology. Online Edition. 17 (16): 1127-1138. doi:10.1080/14772019.2018.1529000.
  23. ^ Barycka, E. (2007). "Evolution and systematics of the feliform Carnivora". Mammalian Biology. 72 (5): 257-282. doi:10.1016/j.mambio.2006.10.011.

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