Carnivora

Carnivoran Temporal range: 42–0  Ma PreꞒ Ꞓ O S D C P T J K Pg N Middle Eocene-Holocene[1][2]
Various carnivorans, with caniforms to the right
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Carnivoramorpha
Clade:	Carnivoraformes
Order:	Carnivora Bowdich, 1821[3]
Suborders
Caniformia Feliformia
Synonyms
list of synonyms: Caniformes (Zagorodniuk, 2008)[4][5] Carnaria (Haeckel, 1866) Carnassidentia (Wortman, 1901) Carnivores (Cuvier, 1817) Carnivori (Vieq d'Azyr, 1792) Carnivorida (Pearse, 1936)[6][7] Carnivoriformes (Kinman, 1994) Carnivoripedida (Vyalov, 1966)[8] Cynofeliformia (Ginsburg, 1982) Cynosia (Rafinesque, 1815) Digitigrada (Illiger, 1811) Digitigradae (Gray, 1821) Eucarnivora (Mekayev, 2002) Ferae (Linnaeus, 1758) Fissipeda (Blumenbach, 1791) Neocarnivora (Radinsky, 1977) Plantigrada (Illiger, 1811)

Carnivora /kɑːrˈnɪvərə/ is an order of placental mammals that have specialized in primarily eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the fifth largest order of mammals, comprising at least 279 species.

Carnivorans live on every major landmass and in a variety of habitats, ranging from the cold polar regions to the hyper-arid region of the

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. The 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 seals. Members of this group are found worldwide and with immense diversity in their diet, behavior, and morphology.^[9]

Etymology

The word carnivore is derived from Latin carō (stem carn-) 'flesh' and vorāre 'to devour', and refers to any meat-eating organism.

Phylogeny

The oldest known carnivoran line mammals (

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.^[12][13] 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.^[14] 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,^[15] 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,^[16] and this suggests a separate evolutionary history and an order-level distinction.^[17] 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.^[14]

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, genet-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 carnivoramorphs that, while viverravids are monophyletic basal group, the miacids are paraphyletic in respect to Carnivora (as shown in the phylogeny below).^{[18][19][20][21][22][23][24][25]}

Pan-Carnivora	†  †Hyaenodonta  †Altacreodus †Tinerhodon  ?  †Wyolestidae  ?  †Simidectes †  sensu lato   Carnivoramorpha   †Viverravidae  †Viverravoidea  ?  †Ravenictis  ?  †Carnivoramorpha sp. (UALVP 50993 & UALVP 50994)  sensu lato   ?  †Carnivoramorpha sp. (UALVP 31176)  ?  †Carnivoramorpha sp. (USNM 538395)  ?  †"Sinopa" insectivorus  Carnivoraformes   Clade "B"  †Eogale †"Miacis" sp. (CM 67873 & CM 77299) †Gracilocyon igniculus †Gracilocyon winkleri †Gracilocyon solei †Gracilocyon †Gracilocyon sp. (Jibou, Romania) †Gracilocyon rosei †Gracilocyon rundlei †Paramiacis †Paroodectes  †Oodectes jepseni   †Oodectes   †Oodectes herpestoides   †Oodectes proximus   †Oodectes pugnax   †Uintacyon hookeri   †Uintacyon rudis   †Uintacyon   †Uintacyon massetericus   †Uintacyon vorax   †Uintacyon asodes   †Uintacyon edax   †Uintacyon acutus   †Uintacyon jugulans   †Uintacyon gingerichi  †Xinyuictis †Zodiocyon †Miocyon †Simamphicyon  †Uintacyon sp. (USGS 1983)  †Messelogale †Quercygalidae  Clade "C"  †Palaearctonyx  († Phlaodectes )  † Vulpavus ovatus †Vulpavus profectus †Vulpavus australis †Vulpavus canavus †Vulpavus completus †Vulpavus †Vulpavus farsonensis †Vulpavus palustris †Vassacyon † †"Miacis" exiguus †"Miacis" deutschi †Africtis  ?  †Carnivoraformes undet. Genus B †Dawsonicyon †"Miacis" boqinghensis †"Miacis" hookwayi †"Miacis" latidens †"Miacis" petilus  ?  †Carnivoraformes undet. Genus A †  Clade "D"  †Lycarion †"Miacis" hargeri  ?  †Ceruttia †"Miacis" invictus †"Miacis" lushiensis †Neovulpavus †Harpalodon †Procynodictis †Prodaphaenus  ?  †Walshius †"Miacis" gracilis †Tapocyon Carnivora (sensu stricto)  (Carnivora [sensu lato])	†Altacreodus/Tinerhodon clade †Gracilocyon/Oodectes clade †Vulpavus clade

Carnivoramorpha as a whole first appeared in the

The phylogenetic relationships of the carnivorans are shown in the following cladogram:[29]^{[30][31][32][33]}

Carnivora

Feliformia   Aeluroidea   Viverroidea   Herpestoidea   Hyaenidae  † Hyaenidae (hyaenas) †Lophocyonidae  sensu lato   Herpestidae  Herpestidae (mongooses)  sensu lato  †Shandgolictis †Asiavorator †Alagtsavbaatar †Anictis   Feloidea    Prionodontidae  Prionodontidae (Asiatic linsangs) †Palaeoprionodon  sensu lato  †Haplogale †Stenoplesictis  ?  †Pseudictis  Felidae  †Barbourofelidae †Viretictis †Stenogale  sensu lato  Nandiniidae (African palm civet) † † Palaeogalidae  Caniformia   † Amphicyonoidea   † †Lycophocyon   Canoidea     Cynoidea    Arctoidea   Ursida    Ursoidea   Ursidae (bears) †Adracon   Mustelida    Musteloidea   ?  †Peignictis †Plesiogale   Pan-Pinnipedia   †Semantoridae  Pinnipedimorpha  † Enaliarctidae †Pinnarctidion   Pinnipediformes   †Pacificotaria †Pteronarctos   Pinnipedia    Otarioidea  Otariidae (eared seals) Odobenidae (walruses)   Phocoidea   †Desmatophocidae Phocidae (earless seals)  sensu stricto   (Pinnipedia sensu lato)

Classification of the extant carnivorans

In 1758 the

• 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 paraphyletic in respect to Pinnipedia,

• Mongooses and a handful of Malagasy endemic species are found to be in a clade with hyenas, with the Malagasy species being in their own family Eupleridae.^[43][44][45]
• The African palm civet is a basal cat-like carnivoran.^[46]
• The linsang is more closely related to cats.^[47]
• Pandas are not procyonids nor are they a natural grouping.^[48] The giant panda is a true bear^[49][50] while the red panda is a distinct family.^[51]
• Skunks and stink badgers are placed in their own family, and are the sister group to a clade containing Ailuridae, Procyonidae and Mustelidae sensu stricto.^[52][51]

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^[53] and 2014^[54] respectively:

Carnivora Bowdich, 1821
Feliformia Kretzoi, 1945
Nandinioidea Pocock, 1929
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Nandiniidae Pocock, 1929	African Palm Civet	Sub-Saharan Africa	1	Nandinia binotata (J. E. Gray, 1830)
Feloidea G. Fischer de Waldheim, 1817
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Felidae G. Fischer de Waldheim, 1817	Cats	Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands)	37	Felis catus Linnaeus, 1758
Prionodontidae Horsfield, 1822	Linsangs	Indomalayan realm	2	Prionodon linsang (Hardwicke, 1821)
Viverroidea J. E. Gray, 1821
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Viverridae J. E. Gray, 1821	Civets, genets, and oyans	Southern Europe, Indomalayan realm, and Africa (introduced to Madagascar)	34	Viverra zibetha Linnaeus, 1758
Herpestoidea Bonaparte, 1845
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Hyaenidae J. E. Gray, 1821	Hyenas	Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent	4	Hyaena hyaena (Linnaeus, 1758)
Herpestidae Bonaparte, 1845	Mongooses	Iberian Peninsula, Africa, the Middle East, the Caucasus, Central Asia, and the Indomalayan realm	34	Herpestes ichneumon (Linnaeus, 1758)
Eupleridae Chenu, 1850	Malagasy mongooses and civets	Madagascar	8	Eupleres goudotii Doyère, 1835
Caniformia Kretzoi, 1945
Canoidea G. Fischer de Waldheim, 1817
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Canidae G. Fischer de Waldheim, 1817	Dogs	Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands)	35	Canis familiaris Linnaeus, 1758
Ursoidea G. Fischer de Waldheim, 1817
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Ursidae G. Fischer de Waldheim, 1817	Bears	Americas and Eurasia	8	Ursus arctos Linnaeus, 1758
Phocoidea J. E. Gray, 1821
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Odobenidae J. A. Allen, 1880	Walrus	The North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere	1	Odobenus rosmarus (Linnaeus, 1758)
Otariidae J. E. Gray, 1825	Eared Seals	Subpolar, temperate, and equatorial waters throughout the Atlantic Oceans	15	Otaria flavescens (Linnaeus, 1758)
Phocidae J. E. Gray, 1821	Earless Seals	The sea and Lake Baikal	18	Phoca vitulina Linnaeus, 1758
Musteloidea G. Fischer de Waldheim, 1817
Family	English Name	Distribution	Number of Extant Species	Type Taxon	Image Figure
Mephitidae Bonaparte, 1845	Skunks and stink badgers	Americas, western Philippines, and Indonesia and Malaysia	12	Mephitis mephitis (Schreber, 1776)
Ailuridae J. E. Gray, 1843	Red Panda	Eastern southwestern China	1	Ailurus fulgens F. Cuvier, 1825
Procyonidae J. E. Gray, 1825	Raccoons	Americas (introduced to Europe, the Caucasus, and Japan)	12	Procyon lotor (Linnaeus, 1758)
Mustelidae G. Fischer de Waldheim, 1817	Weasels, otters, and badgers	Americas, Africa, and Eurasia (introduced to Australasia and several islands)	57	Mustela erminea Linnaeus, 1758

Anatomy

Skull