Carnivora

Carnivora is an order of placental mammals that have specialized in primarily eating flesh. Its members are formally referred to as carnivorans, though some species are omnivorous, like raccoons and bears, and quite a few species like pandas are specialized herbivores. The word 'carnivore' is derived from Latin carō (stem carn-) "flesh" and vorāre "to devour", it refers to any meat-eating organism. The order Carnivora is the fifth largest order of mammals and one of the more successful members of the group; it comprises at least 279 species living 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. They come in a huge array of different body plans in contrasting shapes and sizes. The smallest carnivoran is the least weasel (Mustela nivalis) with a body length of about 11 cm (4.3 in) and a weight of about 25 g (0.88 oz). The largest is the southern elephant seal (Mirounga leonina), with adult males weighing up to 5,000 kg (11,000 lb) and measuring up to 6.7 m (22 ft). 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.[3][4] 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.

For information on all meat-eating organisms, see Carnivore. For the alternative medicine "Carnivora", see Venus flytrap § In alternative medicine.

Carnivorans Temporal range: 42–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Middle Eocene-Holocene[1]
Various carnivorans, with feliforms to the left, and caniforms to the right
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Carnivoramorpha
Clade:	Carnivoraformes
Order:	Carnivora Bowdich, 1821[2]
Suborders
Caniformia Feliformia

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 viverrids. 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 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.

After primates, 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 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 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 (Neovison macrodon) in 1894; the Japanese Sea Lion (Zalophus japonicus) in 1951 and the Caribbean Monk Seal (Neomonachus tropicalis) in 1952.[3] 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.[5]

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

Ferae

Pholidotamorpha       †Oxyaenodonta      †Hyaenodonta    †Hyaenodonta (sensu stricto)       †Altacreodus     †Tinerhodon      sensu lato   Carnivoramorpha     †Viverravoidea  †Viverravidae    ?  †carnivoramorph sp. (UALVP 50993 & UALVP 50994)    ?  †Ravenictis      ?  †carnivoramorph sp. (UALVP 31176)    ?  †carnivoramorph sp. (WW-84: USNM 538395)    Carnivoraformes      †Neovulpavus    ?  †carnivoraform undet. Genus A (UCMP 110072) Neovulpavus clade     †"Miacis" medius      ?  †carnivoraform undet. Genus B (SDSNH 56335)               †Dormaalocyon     †"Miacis" exiguus         †Vassacyon     †Vulpavus         †"Miacis" deutschi   Vulpavus clade         †Miocyon     †Simamphicyon       †Zodiocyon       †Uintacyon    ?  †"Miacis" lushiensis    ?  †"Miacis" thailandicus    ?  †"Miacis" invictus     Uintacyon clade     †Miacis     †carnivoraform sp. (PM 3868)     †Dawsonicyon     †"Miacis" petilus     †"Miacis" latidens    ?  †"Miacis" boqinghensis    ?  †"Miacis" hookwayi    ?  †"Miacis" vulpinus       †Paroodectes     †Paramiacis     †Eogale       †Gracilocyon     †Oodectes     Oodectes clade      ?  †Ceruttia       †Prodaphaenus    ?  †Walshius    Carnivora    Feliformia     Caniformia    sensu stricto         (Carnivora sensu lato)

Carnivoramorpha as a whole first appeared in the Paleocene of North America about 60 million years ago.[4] 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 representatives 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)     †Ceruttia     †Walshius    Carnivora   Feliformia     Aeluroidea     Viverroidea   Herpestoidea   Hyaenidae    †Percrocutidae     Hyaenidae (hyaenas)     †Lophocyonidae    sensu lato   Herpestidae    Herpestidae (mongoose)     Eupleridae (Malagasy mongooses)    sensu lato      Viverridae (viverrids)           †Shandgolictis     †Asiavorator     †Alagtsavbaatar       †Anictis    Feloidea     Prionodontidae    Prionodontidae (Asiatic linsangs)     †Palaeoprionodon    sensu lato    †Haplogale       †Stenoplesictis    ?  †Pseudictis    Felidae    †Barbourofelidae       †Viretictis       †Stenogale     Felidae (cats)        sensu lato          Nandiniidae (African palm civet)       †Nimravidae (false saber-toothed cats)       †Palaeogalidae      Caniformia     †Amphicyonoidea  †Amphicyonidae ("bear-dogs")         †Lycophocyon    Canoidea   Cynoidea  Canidae (canids)    Arctoidea     Ursida   Ursoidea  Ursidae (bears)     †Adracon          Musteloidea          Procyonidae (raccoons)     Mustelidae (mustelids)       Ailuridae      ?  †Peignictis     Mephitidae (skunks)       †Plesiogale      Pinnipedimorpha    †Semantoridae       †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 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.[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:

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 Pacific and Southern Oceans and the southern Indian and Atlantic Oceans	15	Otaria byronia (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 Himalayas and 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 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 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 (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 digitigrade 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] Male carnivorans usually have bacula, though they are absent in hyenas and binturongs.[51]

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

• List of carnivorans by population
• List of species in order Carnivora

References

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External links

• High-Resolution Images of Carnivore Brains