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Temporal range: 42–0  Ma
Carnivora portraits.jpgFossa
Various carnivorans, with feliforms to the left, and caniforms to the right
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Clade: Carnivoramorpha
Clade: Carnivoraformes
Order: Carnivora
Bowdich, 1821 [3]

Carnivora /kɑːrˈnɪvərə/ is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, and have evolved to specialize in eating flesh. The order 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 Sahara Desert to the open seas. They come in a very large array of different body plans in contrasting shapes and sizes.

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 incredible diversity in their diet, behavior, and morphology.


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


The oldest known carnivoran line mammals (Carnivoramorpha) appeared in North America 6 million years after the Cretaceous–Paleogene extinction event. [4] [5] 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 later the creodonts were occupying the megafaunal faunivorous niche. However, following the extinction of mesonychians and the oxyaenid creodonts at the end of the Eocene, carnivorans quickly moved into this niche, with forms like the nimravids being the dominant large-bodied ambush predators during the Oligocene alongside the hyaenodont creodonts (which similarly produced larger, more open-country forms at the start of the Oligocene). By the time Miocene epoch appeared, most if not all of the major lineages and families of carnivorans had diversified and become the most dominant group of large terrestrial predators in Eurasia and North America, with various lineages being successful in megafaunal faunivorous niches at different intervals during the Miocene and later epochs.



Life reconstruction of Tapocyon robustus, a species of miacid Tapocyon robustus.jpg
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. [6] [7] 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. [8] 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. [9] 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. [10] and this suggests a separate evolutionary history and an order-level distinction. [11] 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. [8]

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). [12] [13] [14] [15] [16] [17] [18] [19]


Pholidotamorpha Pangolin Hardwicke (white background).jpg


Oxyaenodonta Patriofelis ferox by R. B. Horsfall (coloured).png


Hyaenodonta(sensu stricto) Hyaenodon horridus by R. B. Horsfall (coloured).jpg



 sensu lato 



carnivoramorph sp. (UALVP 50993 & UALVP 50994)




carnivoramorph sp. (UALVP 31176)


carnivoramorph sp. (WW-84: USNM 538395)




carnivoraform undet. Genus A (UCMP 110072)


"Miacis" medius


carnivoraform undet. Genus B (SDSNH 56335)




"Miacis" exiguus



"Miacis" deutschi







"Miacis" lushiensis


"Miacis" thailandicus


"Miacis" invictus


Miacis Miacis restoration.jpg

carnivoraform sp. (PM 3868)


"Miacis" petilus

"Miacis" latidens


"Miacis" boqinghensis


"Miacis" hookwayi


"Miacis" vulpinus












Feliformia Stamp-russia2014-save-russian-cats-(snow leopard).png

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

 sensu stricto 
 (Carnivorasensu lato) 

Carnivoramorpha as a whole first appeared in the Paleocene of North America about 60 million years ago. [5] Crown 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 representatives of carnivorans were small as the creodonts (specifically, the oxyaenids) and mesonychians dominated the apex predator niches during the Eocene, but in the Oligocene carnivorans became a dominant group of apex predators with the nimravids, and by the Miocene most of the extant carnivoran families have diversified and become the primary terrestrial predators in the Northern Hemisphere.

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


Miacidae(paraphyletic family) Miacis restoration.jpg




Percrocutidae Dinocrocuta gigantea.jpg

Hyaenidae (hyaenas) Hyaena maculata - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg


 sensu lato 

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

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

 sensu lato 

Viverridae (viverrids) Malay Civet (Viverra tangalunga) white background.jpg






Prionodontidae (Asiatic linsangs) Prionodon maculosus.png


 sensu lato 









Felidae (cats) Stamp-russia2014-save-russian-cats-(snow leopard).png

 sensu lato 

Nandiniidae (African palm civet) The carnivores of West Africa (Nandinia binotata white background).png

Nimravidae (false saber-toothed cats) Dinictis Knight.jpg



Amphicyonidae ("bear-dogs") Ysengrinia.jpg



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


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

Ailuridae RedPandaFullBody white background.JPG

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



Mustelidae (mustelids) Fitch white background.png











(eared seals)

Zalophus californianus J. Smit (white background).jpg


USSR stamp Walrus 1977 (white background).png



(earless seals)

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

 sensu stricto 
 (Pinnipediasensu lato) 

Classification of the extant carnivorans

In 1758 the Swedish botanist Carl Linnaeus placed all carnivorans known at the time into the group Ferae (not to be confused with the modern concept of Ferae which also includes pangolins) in the tenth edition of his book Systema Naturae . He recognized 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 1821 that the English writer and traveler Thomas Edward Bowdich gave the group its modern and accepted name. [3]

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]

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 been found to be polyphyletic:

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:

Carnivora Bowdich, 1821
Feliformia Kretzoi, 1945
Nandinioidea Pocock, 1929
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Nandiniidae Pocock, 1929African Palm Civet Sub-Saharan Africa 1 Nandinia binotata (J. E. Gray, 1830)
Feloidea G. Fischer de Waldheim, 1817
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Felidae G. Fischer de Waldheim, 1817Cats Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands)37 Felis catus Linnaeus, 1758 Felis catus-cat on snow.jpg
Prionodontidae Horsfield, 1822Linsangs Indomalayan realm 2 Prionodon linsang (Hardwicke, 1821)
Viverroidea J. E. Gray, 1821
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Viverridae J. E. Gray, 1821Civets, genets, and oyansSouthern Europe, Indomalayan realm, and Africa (introduced to Madagascar)34 Viverra zibetha Linnaeus, 1758 Large Indian Civet, Viverra zibetha in Kaeng Krachan national park.jpg
Herpestoidea Bonaparte, 1845
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Hyaenidae J. E. Gray, 1821Hyenas Africa, the Middle East, the Caucasus, Central Asia, and the Indian subcontinent 4 Hyaena hyaena (Linnaeus, 1758) Hyena at chattbir zoo.jpg
Herpestidae Bonaparte, 1845Mongooses Iberian Peninsula, Africa, the Middle East, the Caucasus, Central Asia, and the Indomalayan realm 34 Herpestes ichneumon (Linnaeus, 1758) Herpestes ichneumon Egipetskii mangust, ili faraonova krysa, ili ikhnevmon.jpg
Eupleridae Chenu, 1850Malagasy mongooses and civets Madagascar 8 Eupleres goudotii Doyère, 1835 Eupleres goudotii - Museo Civico di Storia Naturale Giacomo Doria - Genoa, Italy - DSC02711.JPG
Caniformia Kretzoi, 1945
Canoidea G. Fischer de Waldheim, 1817
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Canidae G. Fischer de Waldheim, 1817Dogs Americas, Africa, and Eurasia (introduced to Madagascar, Australasia and several islands)35 Canis familiaris Linnaeus, 1758 2013072515020909 MyDogs 622.jpg
Ursoidea G. Fischer de Waldheim, 1817
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Ursidae G. Fischer de Waldheim, 1817Bears Americas and Eurasia 8 Ursus arctos Linnaeus, 1758 Kamchatka Brown Bear near Dvuhyurtochnoe on 2015-07-23.jpg
Phocoidea J. E. Gray, 1821
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Odobenidae J. A. Allen, 1880WalrusThe North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere 1 Odobenus rosmarus (Linnaeus, 1758) Walrus2.jpg
Otariidae J. E. Gray, 1825Eared SealsSubpolar, temperate, and equatorial waters throughout the Pacific and Southern Oceans and the southern Indian and Atlantic Oceans15 Otaria flavescens (Linnaeus, 1758) Southern Sea Lions.jpg
Phocidae J. E. Gray, 1821Earless SealsThe sea and Lake Baikal 18 Phoca vitulina Linnaeus, 1758 White harbor seal on moss by Dave Withrow, NOAA.png
Musteloidea G. Fischer de Waldheim, 1817
FamilyEnglish NameDistributionNumber of Extant SpeciesType TaxonImage Figure
Mephitidae Bonaparte, 1845Skunks and stink badgers Americas, western Philippines, and Indonesia and Malaysia 12 Mephitis mephitis (Schreber, 1776) Skunk about to spray.jpg
Ailuridae J. E. Gray, 1843Red PandaEastern Himalayas and southwestern China 1 Ailurus fulgens F. Cuvier, 1825 RedPanda SingalilaNationalPark DFrame.jpg
Procyonidae J. E. Gray, 1825Raccoons Americas (introduced to Europe, the Caucasus, and Japan)12 Procyon lotor (Linnaeus, 1758) Waschbaer auf dem Dach.jpg
Mustelidae G. Fischer de Waldheim, 1817Weasels, otters, and badgers Americas, Africa, and Eurasia (introduced to Australasia and several islands)57 Mustela erminea Linnaeus, 1758 Stoat - RSPB Sandy (28058976023).jpg



Skull of a fossa (Cryptoprocta ferox). Note the large and conical canine and carnassial teeth common in feliforms. Em - Cryptoprocta ferox - 2.jpg
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 stress resistant. All of the terrestrial species of carnivorans have three incisors on each side of each jaw (the exception is the sea otter (Enhydra lutris) which only has two lower incisor teeth). [49] The third molar has been lost. The carnassial pair is made up of the fourth upper premolar and the first lower molar teeth. Like most mammals, the dentition is heterodont, though in some species, such as 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 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. Often a sagittal crest is present, sometimes more evident in sexually dimorphic species such as sea lions and fur seals, though it has also been greatly reduced in some small carnivorans. [50] The braincase is enlarged with the frontoparietal bone at the front. In most species, the eyes are at the front of the face. In caniforms, the rostrum is usually long with many teeth, while in feliforms it is shorter with fewer teeth. The carnassial teeth of feliforms are generally more sectional[ clarification needed ] than those of caniforms. The turbinates are large and complex in comparison to other mammals, providing a large surface area for olfactory receptors. [50]

Postcranial region

A black-backed jackal (Lupulella 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. Canis mesomelas vs. Arctocephalus pusillus.jpg
A black-backed jackal (Lupulella 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 carnivorans as a group. [49] All species of carnivorans are quadrupedal and most have five digits on the front feet and four digits on the back feet. In terrestrial carnivorans, the feet have soft pads. The feet can either be digitigrade as seen in cats, hyenas and dogs or plantigrade as seen in bears, skunks, raccoons, weasels, civets and mongooses. In pinnipeds, the limbs have been modified into flippers.

Members of the Carnivora order, like this tiger, have pads on their feet. Panthera tigris11.jpg
Members of the Carnivora order, like this tiger, have pads on their feet.

Unlike cetaceans and sirenians, which have fully functional tails to help them swim, pinnipeds use their limbs underwater to swim. Earless seals use their back flippers; sea lions and fur seals use their front flippers, and the walrus use all of their limbs. As a result, pinnipeds have significantly shorter tails than other carnivorans.

Aside from the pinnipeds, dogs, bears, hyenas, and cats all have distinct and recognizable appearances. Dogs are usually cursorial mammals and are gracile in appearance, often relying on their teeth to hold prey; bears are much larger and rely on their physical strength to forage for food. Compared to dogs and bears, cats have longer and stronger forelimbs 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 hind legs. The raccoon family and 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] Male carnivorans usually have bacula, though they are absent in hyenas and binturongs. [51]

The length and density of the fur vary depending on the environment that the species inhabits. In warm climate species, the fur is often short in length and lighter. In cold climate species, the fur is either dense or long, often with an oily substance that helps to retain heat. The pelage coloration differs between species, often including black, white, orange, yellow, red, and many shades of grey and brown. Some are striped, spotted, blotched, banded, or otherwise boldly patterned. There seems to be a correlation between habitat and color pattern; for example spotted or banded species tend to be found in heavily forested environments. [49] Some species like the grey wolf are polymorphic with different individual having different coat colors. The arctic fox (Vulpes lagopus) and the stoat (Mustela erminea) have fur that changes from white and dense in the winter to brown and sparse in the summer. In pinnipeds, polar bears, and sea otters a thick insulating layer of blubber helps maintain their body temperature.

Relationship with humans

Carnivorans are arguably the group of mammals of most 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 taxon. In the last 10,000 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 are popular, and they are often charismatic megafauna. Many civilizations have 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 broadly hunted, resulting in extirpation in some areas. Habitat loss and human encroachment as well as climate change have been the primary cause of many species going into decline. Four species of carnivorans have gone extinct since the 1600s: Falkland Island Wolf (Dusicyon australis) in 1876; the Sea Mink (Neogale macrodon) in 1894; the Japanese Sea Lion (Zalophus japonicus) in 1951 and the Caribbean Monk Seal (Neomonachus tropicalis) in 1952. [4] Some species such as the red fox (Vulpes vulpes) and stoat (Mustela erminea) have been introduced to Australasia and have caused many native species to become endangered or even extinct. [52]

See also

Related Research Articles

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A saber-tooth is any member of various extinct groups of predatory therapsids, predominantly carnivoran mammals, that are characterized by long, curved saber-shaped canine teeth which protruded from the mouth when closed. Saber-toothed mammals have been found almost worldwide from the Eocene epoch to the end of the Pleistocene epoch 42 million years ago (mya) – 11,000 years ago (kya).

<span class="mw-page-title-main">Caniformia</span> Suborder of mammals

Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include dogs, bears, raccoons, and mustelids. The Pinnipedia are also assigned to this group. The center of diversification for the Caniformia is North America and northern Eurasia. Caniformia stands in contrast to the other suborder of Carnivora, the Feliformia, the center of diversification of which was in Africa and southern Asia.

<span class="mw-page-title-main">Nimravidae</span> Extinct family of mammals in the order Carnivora

Nimravidae is an extinct family of carnivorans, sometimes known as false saber-toothed cats, whose fossils are found in North America and Eurasia. Not considered to belong to the true cats, the nimravids are generally considered closely related and classified as a distinct family in the suborder Feliformia. Fossils have been dated from the Middle Eocene through the Late Miocene epochs, spanning about 33.2 million years.

<span class="mw-page-title-main">Creodonta</span>

Creodonta is an former order of extinct flesh-eating placental mammals that lived from the early Paleocene to the late Miocene epochs in North America, Europe, Asia and Africa. Originally thought to be a single group of animals ancestral to the modern Carnivora, this order is now usually considered a polyphyletic assemblage of two different groups, the Oxyaenids and the Hyenodonts, not a natural group. Oxyaenids are first known from the Palaeocene of North America, while hyaenodonts hail from the Palaeocene of Africa.

<span class="mw-page-title-main">Mesonychid</span> Extinct taxon of carnivorous ungulates

Mesonychia is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychids first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely when the last genus, Mongolestes, became extinct in the early Oligocene. In Asia, the record of their history suggests they grew gradually larger and more predatory over time, then shifted to scavenging and bone-crushing lifestyles before the group became extinct.

<span class="mw-page-title-main">Ferae</span> A clade of mammals consisting of Carnivores and Pholidotes

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<span class="mw-page-title-main">Carnassial</span> Mammal tooth type

Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified fourth upper premolar and the first lower molar, however this may vary in family. These teeth are also referred to as sectorial teeth.

<span class="mw-page-title-main">Viverravidae</span> Extinct family of carnivores

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<span class="mw-page-title-main">Miacidae</span> Extinct family of carnivores

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<span class="mw-page-title-main">Feliformia</span> Suborder of carnivores

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<span class="mw-page-title-main">Arctoidea</span> Infraorder of mammals

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Hyaenodonta is an extinct order of hypercarnivorous placental pan-carnivoran mammals from mirorder Ferae. Hyaenodonts were important mammalian predators that arose during the early Paleocene in Europe and persisted well into the late Miocene.

<span class="mw-page-title-main">Amphicynodontidae</span> Extinct clade of mammals

Amphicynodontidae is a probable clade of extinct arctoids. While some researchers consider this group to be an extinct subfamily of bears, a variety of morphological evidence links amphicynodontines with pinnipeds, as the group were semi-aquatic otter-like mammals. In addition to the support of the pinniped–amphicynodontine clade, other morphological and some molecular analyses support bears being the closest living relatives to pinnipeds. According to McKenna and Bell (1997) Amphicynodontinae are classified as stem-pinnipeds in the superfamily Phocoidea. Fossils of these mammals have been found in Europe, North America and Asia. Amphicynodontines should not be confused with Amphicyonids (bear-dogs), a separate family of Carnivora which is a sister clade to arctoids within the caniforms, but which may be listed as a clade of extinct arctoids in older publications.

<span class="mw-page-title-main">Herpestoidea</span> Superfamily of mammals

Herpestoidea is a superfamily of mammalia carnivores which includes mongooses, Malagasy carnivorans and the hyenas.

<span class="mw-page-title-main">Carnivoraformes</span> Clade of placental mammals

Carnivoraformes is a clade of placental mammals that includes the modern order Carnivora and its extinct stem-relatives.


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