Caniformia

Caniformia, or Canoidea (literally "dog-like"), is a suborder within the order Carnivora. They typically possess a long snout and nonretractile claws (in contrast to the cat-like carnivorans, the Feliformia). The Pinnipedia (seals, walruses and sea lions) are also assigned to this group. The center of diversification for Caniformia is North America and northern Eurasia. This contrasts with the feliforms, the center of diversification of which was in Africa and southern Asia.

Caniforms
Temporal range: 42–0 Ma
Eocene-Holocene
Caniform portraits
All extant caniform families (clockwise from top left): Canidae, Ursidae, Procyonidae, Mustelidae, Otariidae, Phocidae, Odobenidae, Mephitidae, Ailuridae
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Kretzoi, 1943
Families

 Amphicyonidae† ("bear-dogs")
 Canidae (wolves, foxes)
 Hemicyonidae† ("dog-bears")
 Ursidae (bears)
 Ailuridae (red pandas)
 Enaliarctidae† (extinct pinnipeds)
 Odobenidae (walrus)
 Otariidae (sea lions)
 Phocidae (true seals)
 Mephitidae (skunks)
 Mustelidae (weasels, wolverines)
 Procyonidae (raccoons, kinkajous)

Description

Most members of this group have nonretractile claws (the fisher,[1] marten,[2] red panda,[3] and ringtail have retractile or semiretractile claws[4]) and tend to be plantigrade (with the exception of the Canidae). Other traits that separate Caniformia from Feliformia is that caniforms have longer jaws and have more teeth, with less specialized carnassial teeth. They also tend more towards omnivory and opportunistic feeding, while the feliforms, other than the viverrids, are more specialized for eating meat. Caniforms have single-chambered or partially divided auditory bullae, composed of a single bone, while in feliforms, the auditory bullae are double-chambered, composed of two bones joined by a septum.[5] In the Canoidea, the bulbourethral glands and vesicula seminalis are always absent. Relative to body size, the baculum is usually longer in the Canoidea than in the Feloidea.[5]

Extant families

Polar-bear-arctic-wildlife-snow-53425
Polar bear, the largest terrestrial caniform
Mustela nivalis (two, fighting)
The smallest caniform is the least weasel.

Caniformia consists of nine extant families, with three extinct families also recognized. The extant families are monophyletic according to phylogenetic molecular analysis.[6] At one time, the Hyaenidae (hyenas) were included, but genetic testing has shown them to belong in Feliformia, instead. Terrestrial caniforms in the wild are found on all continents with the exception of Antarctica, while pinnipeds are distributed throughout the world's oceans.

Family Canidae (dogs and other canids) includes wolves, dogs, coyotes, and foxes, as well as a number of less familiar animals. The family is currently divided into two major groups, the true dogs (tribe Canini), which includes nine genera, and the true foxes (tribe Vulpini) with two genera. In addition, two basal genera are described. About 35 species of extant canids are currently recognized. Canids are the most social of all caniforms, sometimes living in packs. The dog is the most diverse of all mammals in terms of body structure variants.

Family Ursidae (bears) is the largest of all the land caniforms. Eight species are recognized, divided into five genera. They range from the large polar bear (males, 350–680+ kg or 775-1500+ lb) to the small sun bear (males, 30–60 kg or 66–132 lb) and from the endangered giant panda to the very common black bear. Common characteristics of modern bears include a large body with stocky legs, a long snout, shaggy hair, plantigrade paws with five nonretractile claws, and a short tail. Most bears are omnivorous, with largely varied diets that include both plants and animals. The polar bear is the most carnivorous of bears due to the arctic climate in which it lives, and shows a preference for eating seals. The giant panda is the most herbivorous bear and has evolved a number of adaptations, including a sixth "toe", specialized teeth, and strong jaw muscles, to allow it to feed nearly exclusively on bamboo, a tough member of the grass family. The sloth bear has some adaptations for ant and termite eating, with a long snout, powerful claws, and missing upper front teeth, though it also eats honey and fruit.

Family Ailuridae consists today of a single species, the red panda, which was once thought to be included in the Procyonidae or Ursidae lineages, but is now placed in its own family along with a number of extinct species. It is found in the Himalayas, including southern China, Nepal, Bhutan, India, and Pakistan. Fossil species of the family are also found in North America.[7]

Family Mephitidae (skunks and stink badgers) was once classified as mustelids, but are now recognized as a lineage in their own right. The 12 species of skunks are divided into four genera: Mephitis (hooded and striped skunks, two species), Spilogale (spotted skunks, four species), Mydaus (stink badgers, two species) and Conepatus (hog-nosed skunks, four species). The two skunk species in the genus Mydaus inhabit Indonesia and the Philippines; all other skunks inhabit the Americas from Canada to central South America.

Family Mustelidae (badgers, weasels and otters) is the largest family of carnivora, with 22 extant genera and roughly 57 extant species. While highly variable in shape, size, and behavior, most mustelids are smaller animals with short legs, short, round ears, and thick fur. Mustelids are predominantly carnivorous. While not all share identical dentition, they all possess teeth adapted for eating flesh, including the presence of shearing carnassials.

Members of Family Procyonidae (raccoons, coatis) are smallish animals, with generally slender bodies and long tails. Nineteen extant species in six genera are currently recognized. Except for the kinkajou, all procyonids have banded tails and distinct facial markings, and like bears, are plantigrade, walking on the soles of their feet. Most species have nonretractile claws. Early procyonids may have been an offshoot of the canids that adapted to more omnivorous diets.[8]

Pinnipedia (seals, sea lions, and walruses clade) is a widely distributed and diverse group of semiaquatic marine mammals which is closely related to an extinct group of pinnipeds, Enaliarctos. While support for the monophyly of pinnipeds is strong, the relationship of pinnipeds to terrestrial mammals is still unclear. Some studies support the hypothesis that the bears are their closest relatives,[9][10][11] while others support a closer relationship to the mustelids.[12][13][14][15]

Pinnipeds split from other caniforms 50 million years ago (mya) during the Eocene.[14]

The clade is currently divided into three families:

Family Phocidae (true or earless seals) consists of around 19 species of highly aquatic, barrel-shaped animals ranging from 45 kg (100 lb) and 1.2 m (4 ft) in length (the ringed seal), to 2,400 kg (5,300 lb) and 5 m (16 ft) (southern elephant seal). Phocids are found throughout the world's oceans.
Family Otariidae (eared seals, sea lions, fur seals) is distributed throughout the world's oceans with the exception of the North Atlantic. The 15 species (divided into seven genera) of otariids are distinguished from phocids by visible external ears (pinnae), more dog-like faces, and the ability to turn their rear flippers forward.
Family Odobenidae currently includes a single species, the walrus. A large (2,000 kg or 4,400 lb), distinctive pinniped with long whiskers and tusks, the walrus has a discontinuous circumpolar distribution in the Arctic Ocean and sub-Arctic seas of the Northern Hemisphere. It is primarily a benthic forager of bivalve mollusks and other marine invertebrates.
Miacis
Miacis is the earliest known member of the order Carnivora.

Evolution

Caniforms first appeared as tree-climbing, superficially marten-like carnivores in the Eocene around 42 million years ago. Miacis cognitus was probably an early caniform. Like many other early carnivorans, it was well suited for tree climbing with needle-sharp claws, and had limbs and joints that resemble those of modern carnivorans. M. cognitus was probably a very agile forest dweller that preyed on smaller animals, such as small mammals, reptiles, and birds.

New zealand sea lion nursing
New Zealand sea lion (Phocarctos hookeri)

Debate continues on the origin of pinnipeds. Recent molecular evidence suggests pinnipeds evolved from a bear-like ancestor about 23 million years ago during the late Oligocene or early Miocene epochs, a transitional period between the warmer Paleogene and cooler Neogene periods.[13] However, discovery of the fossil Puijila darwini in early Miocene deposits in Nunavut suggests a different scenario. Like a modern otter, Puijila had a long tail, short limbs, and webbed feet instead of flippers. However, its limbs and shoulders were more robust, and Puijila likely had been a quadrupedal swimmer–retaining a form of aquatic locomotion that give rise to the major swimming types employed by modern pinnipeds. Puijila has been assigned to a clade of mustelids.

Phylogeny

Procyon lotor (Common raccoon)
Common raccoon (Procyon lotor)

The cladogram is based on molecular phylogeny of six genes in Flynn, 2005.[13]

   Caniformia   

AmphicyonidaeYsengrinia americana

Canidae African golden wolf

   Arctoidea   
   Ursoidea   

HemicyonidaeHemicyon sansaniensis

Ursidae American black bear

Pinnipedia

EnaliarctidaeEnaliarctos mealsi.JPG

   

Phocidae Common seal

   

Otariidae California sea lion

Odobenidae Pacific walrus

   Musteloidea   
   

Ailuridae Red panda

   

Mephitidae Striped skunk

   

Procyonidae Common raccoon

Mustelidae European polecat

References

  1. ^ Rhines, C (2003). "Martes pennanti (On-line)". Animal Diversity Web. Retrieved March 12, 2011.
  2. ^ "American Marten". New York State Department of Environmental Conservation. Retrieved March 2011. Check date values in: |accessdate= (help)
  3. ^ Roberts, M. S.; Gittleman, J. L. (1984). "Ailurus fulgens" (PDF). Mammalian Species. The American Society of Mammalogists (222): 3. doi:10.2307/3503840. Retrieved March 2011. Check date values in: |accessdate= (help)
  4. ^ Goldberg, J. (2003). "Bassariscus astutus (On-line)". Animal Diversity Web. Retrieved March 2011. Check date values in: |accessdate= (help)
  5. ^ a b R. F. Ewer (1973). The Carnivores. Cornell University Press. ISBN 978-0-8014-8493-3. Retrieved 9 January 2013.
  6. ^ Eizirik E.; Murphy W.J.; Koepfli K.P.; Johnson W.E.; Dragoo J.W.; O'Brien S.J. (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. ^ "Two new carnivores from an unusual late Tertiary forest biota in eastern North America" (PDF). Archived from the original (PDF) on 2007-09-30. Retrieved 2007-02-27.
  8. ^ Russell, James (1984). Macdonald, D., ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 98–99. ISBN 0-87196-871-1.
  9. ^ Lento, G. M.; Hickson, R. E.; Chambers, G. K.; Penny, D. (1995). "Use of spectral analysis to test hypotheses on the origin of pinnipeds". Molecular Biology and Evolution. 12 (1): 28–52. doi:10.1093/oxfordjournals.molbev.a040189. PMID 7877495.
  10. ^ Hunt, R. M. Jr.; Barnes, L. G. (1994). "Basicranial evidence for ursid affinity of the oldest pinnipeds" (PDF). Proceedings of the San Diego Society of Natural History. 29: 57–67.
  11. ^ Higdon, J. W.; Bininda-Emonds, O. R.; Beck, R. M.; Ferguson, S. H. (2007). "Phylogeny and divergence of the pinnipeds (Carnivora: Mammalia) assessed using a multigene dataset". BMC Evolutionary Biology. 7: 216. doi:10.1186/1471-2148-7-216. PMC 2245807. PMID 17996107.
  12. ^ Sato, J. J.; Wolsan, M.; Suzuki, H.; Hosoda, T.; Yamaguchi, Y.; Hiyama, K.; Kobayashi, M.; Minami, S. (2006). "Evidence from nuclear DNA sequences sheds light on the phylogenetic relationships of Pinnipedia: Single origin with affinity to Musteloidea". Zoological Science. 23 (2): 125–46. doi:10.2108/zsj.23.125. PMID 16603806.
  13. ^ a b c Flynn, J. J.; Finarelli, J. A.; Zehr, S.; Hsu, J.; Nedbal, M. A. (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.
  14. ^ a b Hammond, J. A.; Hauton, C.; Bennett, K. A.; Hall, A. J. (2012). Nikolaidis, Nikolas, ed. "Phocid seal leptin: Tertiary structure and hydrophobic receptor binding site preservation during distinct leptin gene evolution". PLoS ONE. 7 (4): e35395. doi:10.1371/journal.pone.0035395. PMC 3334926. PMID 22536379.
  15. ^ Rybczynski, N.; Dawson, M. R.; Tedford, R. H. (2009). "A semi-aquatic Arctic mammalian carnivore from the Miocene epoch and origin of Pinnipedia". Nature. 458 (7241): 1021–24. doi:10.1038/nature07985. PMID 19396145.

External links

Amphicyanis

Amphicyanis is an extinct genus of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog"), and which inhabited Eurasia and North America.Amphicyanis was named by Springhorn (1977). It was assigned to Amphicyonidae by Carroll (1988).

Arctoidea

Arctoidea is an infraorder of mostly carnivorous mammals which include the extinct Hemicyonidae (dog-bears), and the extant Musteloidea (weasels, raccoons, skunks, red pandas), Pinnipedia (seals, sea lions), and Ursidae (bears), found in all continents from the Eocene, 46 million years ago, to the present.

Arctoids are caniforms, along with dogs (canids) and extinct bear dogs (Amphicyonidae).

The earliest caniforms were superficially similar to martens, which are tree-dwelling mustelids.

Together with feliforms, caniforms comprise the order Carnivora.

Bear dog

Amphicyonidae is an extinct family of large terrestrial carnivorans belonging to the suborder Caniformia which inhabited North America, Europe, Asia, and Africa from the Middle Eocene subepoch to the Pliocene epoch 42—2.6 Mya, existing for about 39.4 million years. Amphicyonids are often colloquially referred to as "bear-dogs". They are closely related to true dogs (Canidae) and a little less related to bears (Ursidae).

Brachycyon

Brachycyon is an extinct genus of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog"), which inhabited Eurasia and North America. Brachycyon was named by Filhol (1872). It was assigned to Amphicyonidae by Carroll (1988).

Brachyrhynchocyon

Brachyrhynchocyon is an extinct genus of terrestrial carnivore, which belonged to the family Amphicyonidae ("bear dogs") of the suborder Caniformia.

Carnivora

Carnivora (; from Latin carō (stem carn-) "flesh" and vorāre "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").

Carnivoramorpha

Carnivoramorpha are a clade of mammals that includes the modern order Carnivora.

Cynelos

Cynelos is a large extinct genus of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog"), and which inhabited North America, Asia, and Africa from the Early Miocene subepoch to the Late Miocene subepoch 23.3—7.2 Mya, existing for approximately 16.1 million years.Dentition of this animal displayed only the characteristics of a carnivore.

Daphoenodon

Daphoenodon is an extinct genus of terrestrial carnivore, which belonged to the family Amphicyonidae ("bear dogs") of the suborder Caniformia.

Feliformia

Feliformia (also Feloidea) is a suborder within the order Carnivora consisting of "cat-like" carnivorans, including cats (large and small), hyenas, mongooses, civets, and related taxa. Feliformia stands in contrast to the other suborder of Carnivora, Caniformia ("dog-like" carnivorans).

The separation of the Carnivora into the broad groups of feliforms and caniforms is widely accepted, as is the definition of Feliformia and Caniformia as suborders (sometimes superfamilies). The classification of feliforms as part of the Feliformia suborder or under separate groupings continues to evolve.

Systematic classifications dealing with only extant taxa include all feliforms into the Feliformia suborder, though variations exist in the definition and grouping of families and genera. Indeed, molecular phylogenies suggest that all extant Feliformia are monophyletic.The extant families as reflected in the taxa chart at right and the discussions in this article reflect the most contemporary and well-supported views (as at the time of writing this article).

Systematic classifications dealing with both extant and extinct taxa vary more widely. Some separate the feliforms (extant and extinct) as: Aeluroidea (superfamily) and Feliformia (suborder). Others include all feliforms (extant, extinct and "possible ancestors") into the Feliformia suborder. Some studies suggest this inclusion of "possible ancestors" into Feliformia (or even Carnivora) may be spurious. The extinct (†) families as reflected in the taxa chart are the least problematic in terms of their relationship with extant feliforms (with the most problematic being Nimravidae).

Haplocyonoides

Haplocyonoides is an extinct genus of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog"), and which inhabited Europe from the Early Miocene subepoch (20 Mya)—(16.9 Mya). Haplocyonoides existed for approximately 3.1 million years.Haplocyonoides was named by Hürzeler (1940) and was assigned to Amphicyonidae by Carroll (1988).

Haplocyonopsis

Haplocyonopsis is an extinct genus of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog").Lived in Miocene epoch in Europe.

Haplocyonopsis was named by de Bonis (1973) and was assigned to Amphicyonidae by Carroll (1988).

Mephitis (genus)

The genus Mephitis is one of several genera of skunks, which has two species and a North American distribution.

Mustelidae

The Mustelidae (; from Latin mustela, weasel) are a family of carnivorous mammals, including weasels, badgers, otters, ferrets, martens, mink, and wolverines, among others. Mustelids are diverse and the largest family in the order Carnivora, suborder Caniformia. Mustelidae comprises about 56-60 species across eight subfamilies.

Paradaphoenus

Paradaphoenus is a physically small member of the extinct family of terrestrial carnivores belonging to the suborder Caniformia, family Amphicyonidae ("bear dog"), and which inhabited North America from the Early Oligocene subepoch to the Middle Miocene subepoch living 33.3—15.97 Ma, existing for approximately 17.33 million years.

Polecat

Polecat is a common name for mammals in the order Carnivora and subfamilies Galictinae and Mustelinae. Polecats do not form a single taxonomic rank (i.e., clade); the name is applied to several species with broad similarities (including having a dark mask-like marking across the face) to European polecats, the only species native to the British Isles.

In the United States, the term polecat is sometimes applied to the black-footed ferret, a native member of the Mustelinae, and (loosely) to skunks, which are only distantly related.

Despite the name, polecats, being various caniform mustelids, are more closely related to dogs than cats, which is why they belong to the suborder Caniformia.

In Canada, the term polecat is sometimes applied to electric utility linemen.

Pseudamphicyon

Pseudamphicyon is a member of the extinct family Amphicyonidae of terrestrial carnivores belonging to the suborder Caniformia.Pseudamphicyon was named Schlosser in 1899 and was assigned to Amphicyonidae by Carroll (1988).

South American coati

The South American coati or ring-tailed coati (Nasua nasua) is a species of coati and a member of the raccoon family (Procyonidae), from tropical and subtropical South America. In Brazilian Portuguese, it is known as quati. An adult generally weighs 2–7.2 kg (4.4–15.9 lb) and is 85–113 cm (33–44 in) long, with half of that being its tail. Its color is highly variable and the rings on the tail may be only somewhat visible, but its distinguishing characteristic is that it lacks the largely white snout (or "nose") of its northern relative, the white-nosed coati.

Western lowland olingo

The western lowland olingo (Bassaricyon medius) is a species of olingo from Central and South America, where it is known from Panama and from Colombia and Ecuador west of the Andes.

Extant Carnivora species

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