Canidae

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Canids
Temporal range:
Ma
CanisCuonLycaonCerdocyonChrysocyonSpeothosVulpesNyctereutesOtocyonUrocyon
10 of the 13 extant canid genera
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Infraorder: Cynoidea
Family: Canidae
Fischer de Waldheim, 1817[2]
Type genus
Canis
Linnaeus, 1758
Subfamilies and extant genera

Canidae (

jackals
and other species.

Canids are found on all continents except

domestic animals
.

Taxonomy

See also: List of canids

In the history of the carnivores, the family Canidae is represented by the two extinct subfamilies designated as Hesperocyoninae and Borophaginae, and the extant subfamily Caninae.

mastication. This, together with the development of a distinct entoconid cusp and the broadening of the talonid of the first lower molar, and the corresponding enlargement of the talon of the upper first molar and reduction of its parastyle distinguish these late Cenozoic canids and are the essential differences that identify their clade.[1]
: p6 

The cat-like

Late Pliocene.[1]
: 7 

Phylogenetic relationships

This cladogram shows the phylogenetic position of canids within Caniformia, based on fossil finds:[1]

  Caniformia  
  Arctoidea  

(bears, procyonids, pinnipeds, etc.)

Cynoidea
  †Miacis  

(various †Miacis spp.)

  Canidae  
  †Hesperocyoninae  

(Mesocyon, Enhydrocyon, Hesperocyon, and other basal canines, e.g. Cynodictis)

  †Borophaginae  

(Aelurodon, cynarctins, and others)

  Caninae  

(all modern canids and extinct relatives)

Evolution

See also: Caninae § Phylogenetic relationships
Canis aureus
(Caninae)

The Canidae are a diverse group of some 37 species ranging in size from the maned wolf with its long limbs to the short-legged bush dog. Modern canids inhabit forests, tundra, savannas, and deserts throughout tropical and temperate parts of the world. The evolutionary relationships between the species have been studied in the past using morphological approaches, but more recently, molecular studies have enabled the investigation of phylogenetics relationships. In some species, genetic divergence has been suppressed by the high level of gene flow between different populations and where the species have hybridized, large hybrid zones exist.[8]

Eocene epoch

feliform (cat-like). By 40 Mya, the first identifiable member of the dog family had arisen. Named Prohesperocyon wilsoni, its fossils have been found in southwest Texas. The chief features which identify it as a canid include the loss of the upper third molar (part of a trend toward a more shearing bite), and the structure of the middle ear which has an enlarged bulla (the hollow bony structure protecting the delicate parts of the ear). Prohesperocyon probably had slightly longer limbs than its predecessors, and also had parallel and closely touching toes which differ markedly from the splayed arrangements of the digits in bears.[9]

Canidae soon divided into three subfamilies, each of which diverged during the Eocene:

hypercarnivorous diets that made them prone to extinction.[10]
: Fig. 1 

Oligocene epoch

By the

viverrids in appearance. Hesperocyonines eventually became extinct in the middle Miocene. One of the early Hesperocyonines, the genus Hesperocyon, gave rise to Archaeocyon and Leptocyon. These branches led to the borophagine and canine radiations.[11]

Miocene epoch

Around 8 Mya, the Beringian land bridge allowed members of the genus Eucyon a means to enter Asia from North America and they continued on to colonize Europe.[12]

Pliocene epoch

The

mastication and shearing.[11] Around 5 million years ago, some of the Old World Eucyon evolved into the first members of Canis,[13] In the Pliocene, around 4–5 Mya, Canis lepophagus appeared in North America. This was small and sometimes coyote-like. Others were wolf-like. C. latrans (the coyote) is theorized to descend from C. lepophagus.[14]

The formation of the Isthmus of Panama, about 3 Mya, joined South America to North America, allowing canids to invade South America, where they diversified. However, the last common ancestor of the South American canids lived in North America some 4 Mya and more than one incursion across the new land bridge is likely given the fact that more than one lineage is present in South America. Two North American lineages found in South America are the gray fox (Urocyon cinereoargentus) and the now-extinct dire wolf (Aenocyon dirus). Besides these, there are species endemic to South America: the maned wolf (Chrysocyon brachyurus), the short-eared dog (Atelocynus microtis), the bush dog (Speothos venaticus), the crab-eating fox (Cerdocyon thous), and the South American foxes (Lycalopex spp.). The monophyly of this group has been established by molecular means.[12]

Pleistocene epoch

During the

Canis ambrusteri emerged in North America. A large wolf, it was found all over North and Central America and was eventually supplanted by the dire wolf, which then spread into South America during the Late Pleistocene.[15]

By 0.3 Mya, a number of subspecies of the gray wolf (C. lupus) had developed and had spread throughout Europe and northern Asia.[16] The gray wolf colonized North America during the late Rancholabrean era across the Bering land bridge, with at least three separate invasions, with each one consisting of one or more different Eurasian gray wolf clades.[17] MtDNA studies have shown that there are at least four extant C. lupus lineages.[18] The dire wolf shared its habitat with the gray wolf, but became extinct in a large-scale extinction event that occurred around 11,500 years ago. It may have been more of a scavenger than a hunter; its molars appear to be adapted for crushing bones and it may have gone extinct as a result of the extinction of the large herbivorous animals on whose carcasses it relied.[15]

In 2015, a study of mitochondrial genome sequences and whole-genome nuclear sequences of African and Eurasian canids indicated that extant wolf-like canids have colonized Africa from Eurasia at least five times throughout the Pliocene and Pleistocene, which is consistent with fossil evidence suggesting that much of African canid fauna diversity resulted from the immigration of Eurasian ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid conditions. When comparing the African and Eurasian golden jackals, the study concluded that the African specimens represented a distinct monophyletic lineage that should be recognized as a separate species, Canis anthus (

Mya, but the African golden wolf separated 1.3 Mya. Mitochondrial genome sequences indicated the Ethiopian wolf diverged from the wolf/coyote lineage slightly prior to that.[19]
: S1 

Characteristics

Skeleton of a black-backed jackal (Lupulella mesomelas) on display at the Museum of Osteology
Diagram of a wolf skull with key features labelled
Eurasian wolf skull

Wild canids are found on every continent except Antarctica, and inhabit a wide range of different habitats, including

raccoon dog habitually climb trees.[23][24][25]

All canids have a similar basic form, as exemplified by the gray wolf, although the relative length of muzzle, limbs, ears, and tail vary considerably between species. With the exceptions of the bush dog, the raccoon dog and some domestic

mammae.[27]

All canids are digitigrade, meaning they walk on their toes. The tip of the nose is always naked, as are the cushioned pads on the soles of the feet. These latter consist of a single pad behind the tip of each toe and a more-or-less three-lobed central pad under the roots of the digits. Hairs grow between the pads and in the Arctic fox the sole of the foot is densely covered with hair at some times of the year. With the exception of the four-toed African wild dog (Lycaon pictus), five toes are on the forefeet, but the pollex (thumb) is reduced and does not reach the ground. On the hind feet are four toes, but in some domestic dogs, a fifth vestigial toe, known as a dewclaw, is sometimes present, but has no anatomical connection to the rest of the foot. In some species, slightly curved nails are non-retractile and more-or-less blunt[26] while other species have sharper, partially-retractile claws.[citation needed]

The

ungulates used to maintain the posture of the head and neck with little active muscle exertion; this ligament allows them to conserve energy while running long distances following scent trails with their nose to the ground. However, based on skeletal details of the neck, at least some of the Borophaginae (such as Aelurodon) are believed to have lacked this ligament.[30]

Dentition

carnassials that are used together in a scissor-like action to shear the muscle and tendon of prey.[31]

Canids use their premolars for cutting and crushing except for the upper fourth premolar P4 (the upper carnassial) that is only used for cutting. They use their molars for grinding except for the lower first molar m1 (the lower carnassial) that has evolved for both cutting and grinding depending on the canid's dietary adaptation. On the lower carnassial, the

talonid is used for grinding. The ratio between the trigonid and the talonid indicates a carnivore's dietary habits, with a larger trigonid indicating a hypercarnivore and a larger talonid indicating a more omnivorous diet.[32][33] Because of its low variability, the length of the lower carnassial is used to provide an estimate of a carnivore's body size.[32]

A study of the estimated bite force at the canine teeth of a large sample of living and fossil mammalian predators, when adjusted for their body mass, found that for

placental mammals the bite force at the canines was greatest in the extinct dire wolf (163), followed among the modern canids by the four hypercarnivores that often prey on animals larger than themselves: the African wild dog (142), the gray wolf (136), the dhole (112), and the dingo (108). The bite force at the carnassials showed a similar trend to the canines. A predator's largest prey size is strongly influenced by its biomechanical limits.[34]

Most canids have 42

teeth, with a dental formula of: 3.1.4.23.1.4.3. The bush dog has only one upper molar with two below, the dhole has two above and two below. and the bat-eared fox has three or four upper molars and four lower ones.[26] The molar teeth are strong in most species, allowing the animals to crack open bone to reach the marrow. The deciduous, or baby teeth, formula in canids is 3.1.33.1.3, molars being completely absent.[26]

Life history

Social behavior

Dholes attacking a sambar, Bandipur National Park
Sri Lankan jackals grooming each other
See also: Wolf § Behaviour, Dog behavior, Red fox § Behaviour, and African wild dog § Behaviour

Almost all canids are social animals and live together in groups. In general, they are territorial or have a home range and sleep in the open, using their dens only for breeding and sometimes in bad weather.[35] In most foxes, and in many of the true dogs, a male and female pair work together to hunt and to raise their young. Gray wolves and some of the other larger canids live in larger groups called packs. African wild dogs have packs which may consist of 20 to 40 animals and packs of fewer than about seven individuals may be incapable of successful reproduction.[36] Hunting in packs has the advantage that larger prey items can be tackled. Some species form packs or live in small family groups depending on the circumstances, including the type of available food. In most species, some individuals live on their own. Within a canid pack, there is a system of dominance so that the strongest, most experienced animals lead the pack. In most cases, the dominant male and female are the only pack members to breed.[37]

Communication

See also: Dog communication and Wolf communication
Red foxes barking in Pinbury Park, Gloucestershire, England.

Canids communicate with each other by

supracaudal glands.[44]


Reproduction

See also: Canine reproduction
Korean wolves and red foxes
A feral dog from Sri Lanka nursing her pups

Canids as a group exhibit several reproductive traits that are uncommon among mammals as a whole. They are typically

maned wolves are induced ovulators.[47]

During the proestral period, increased levels of

photoperiod stimulus breaks down under conditions of artificial lighting.[45] Canids have an oestrus period of 1 to 20 days, lasting one week in most species.[48]

The size of a litter varies, with from one to 16 or more pups being born. The young are born small, blind and helpless and require a long period of parental care. They are kept in a den, most often dug into the ground, for warmth and protection.[26] When the young begin eating solid food, both parents, and often other pack members, bring food back for them from the hunt. This is most often vomited up from the adult's stomach. Where such pack involvement in the feeding of the litter occurs, the breeding success rate is higher than is the case where females split from the group and rear their pups in isolation.[49] Young canids may take a year to mature and learn the skills they need to survive.[50] In some species, such as the African wild dog, male offspring usually remain in the natal pack, while females disperse as a group and join another small group of the opposite sex to form a new pack.[51]

Canids and humans

fox hunt
Further information: Dog

One canid, the

agriculturists.[58][59]

The fact that wolves are pack animals with cooperative social structures may have been the reason that the relationship developed. Humans benefited from the canid's loyalty, cooperation, teamwork, alertness and tracking abilities, while the wolf may have benefited from the use of weapons to tackle larger prey and the sharing of food. Humans and dogs may have evolved together.[60]

Among canids, only the gray wolf has widely been known to prey on humans.[61][page needed] Nonetheless, at least two records of coyotes killing humans have been published,[62] and at least two other reports of golden jackals killing children.[63] Human beings have trapped and hunted some canid species for their fur and some, especially the gray wolf, the coyote and the red fox, for sport.[64] Canids such as the dhole are now endangered in the wild because of persecution, habitat loss, a depletion of ungulate prey species and transmission of diseases from domestic dogs.[65]

See also

References

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Bibliography
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