Canidae (; from Latin, ''

canis ''Canis'' is a genus of the Caninae which includes multiple extant taxon, extant species, such as Wolf, wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-develo ...

'', " dog") is a biological family of caniform

carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...

ns, constituting a

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

. A member of this family is also called a canid (). The family includes three subfamilies: the

Caninae Caninae (whose members are known as canines () is the only living subfamily within Canidae, alongside the extinct Borophaginae and Hesperocyoninae. They first appeared in North America, during the Oligocene around 35 million years ago, subsequent ...

, and the extinct Borophaginae and Hesperocyoninae. The Caninae are known as canines, and include

domestic dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers ...

wolves The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...

coyote The coyote (''Canis latrans''), also known as the American jackal, prairie wolf, or brush wolf, is a species of canis, canine native to North America. It is smaller than its close relative, the Wolf, gray wolf, and slightly smaller than the c ...

s, raccoon dogs,

foxes Foxes are small-to-medium-sized omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull; upright, triangular ears; a pointed, slightly upturned snout; and a long, bushy tail ("brush"). Twelve species ...

jackal Jackals are Canidae, canids native to Africa and Eurasia. While the word has historically been used for many canines of the subtribe Canina (subtribe), canina, in modern use it most commonly refers to three species: the closely related black-b ...

s and other species. Canids are found on all continents except

Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

, having arrived independently or accompanied by human beings over extended periods of time. Canids vary in size from the gray wolf to the fennec fox. The body forms of canids are similar, typically having long muzzles, upright ears, teeth adapted for cracking bones and slicing flesh, long legs, and bushy tails. They are mostly social animals, living together in family units or small groups and behaving co-operatively. Typically, only the dominant pair in a group breeds and a litter of young are reared annually in an underground den. Canids communicate by scent signals and vocalizations. One canid, the domestic dog, originated from a

symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...

relationship with

Upper Paleolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories ...

humans and is one of the most widely kept domestic animals.

Taxonomy

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. This subfamily includes all living canids and their most recent fossil relatives. All living canids as a group form a dental

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

relationship with the extinct borophagines, with both groups having a

bicuspid The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mou ...

(two points) on the lower

carnassial 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 f ...

talonid, which gives this tooth an additional ability in

mastication Chewing or mastication is the process by which food is comminution, crushed and ground by the teeth. It is the first step in the process of digestion, allowing a greater surface area for digestive enzymes to break down the foods. During the mast ...

. 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 The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

canids and are the essential differences that identify their clade. The cat-like

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

and dog-like

Caniformia Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include Canidae, dogs (Wolf, wolves, foxes, etc.), bears, raccoons, and Mustelidae, mustelids. The Pinnipedia (pinniped, seals, walruses and sea lions) ...

emerged within the Carnivoramorpha around 45–42 Mya (million years ago). The Canidae first appeared in North America during the

Late Eocene The Priabonian is, in the International Commission on Stratigraphy, ICS's geologic timescale, the latest age (geology), age or the upper stage (stratigraphy), stage of the Eocene epoch (geology), Epoch or series (stratigraphy), Series. It spans ...

(37.8-33.9 Mya). They did not reach Eurasia until the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

or to South America until the

Late Pliocene Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...

Phylogenetic relationships

This

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

shows the phylogenetic position of canids within

, based on fossil finds:

Evolution

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 In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

relationships. In some species,

genetic divergence Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes ( mutations) through time, often leading to reproductive isolation and continued mutation even after the populations h ...

has been suppressed by the high level of

gene flow In population genetics, gene flow (also known as migration and allele flow) is the transfer of genetic variation, genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent ...

between different populations and where the species have hybridized, large hybrid zones exist.

Eocene epoch

Carnivora Carnivora ( ) is an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...

ns evolved after the

extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

of the non-avian dinosaurs 66 million years ago. Around 50 million years ago, or earlier, in the

Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

, the Carnivora split into two main divisions:

caniform Caniformia is a suborder within the order Carnivora consisting of "dog-like" carnivorans. They include dogs (wolves, foxes, etc.), bears, raccoons, and mustelids. The Pinnipedia ( seals, walruses and sea lions) are also assigned to this gr ...

(dog-like) and

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

(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

bear Bears are carnivoran mammals of the family (biology), family Ursidae (). They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats ...

s. Canidae soon divided into three subfamilies, each of which diverged during the Eocene: Hesperocyoninae (about 39.74–15 Mya), Borophaginae (about 34–32 Mya), and

(about 34–30 Mya; the only surviving subfamily). Members of each subfamily showed an increase in body mass with time and some exhibited specialized hypercarnivorous diets that made them prone to extinction.

Oligocene epoch

By the

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

, all three subfamilies (Hesperocyoninae, Borophaginae, and Caninae) had appeared in the fossil record of North America. The earliest and most primitive branch of the Canidae was Hesperocyoninae, which included the coyote-sized '' Mesocyon'' of the Oligocene (38–24 Mya). These early canids probably evolved for the fast pursuit of prey in a grassland habitat; they resembled modern 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.Martin, L.D. 1989. Fossil history of the terrestrial carnivora. Pages 536–568 in J.L. Gittleman, editor. Carnivore Behavior, Ecology, and Evolution, Vol. 1. Comstock Publishing Associates: Ithaca.

Miocene epoch

Around 8 Mya, the

Beringia Beringia is defined today as the land and maritime area bounded on the west by the Lena River in Russia; on the east by the Mackenzie River in Canada; on the north by 70th parallel north, 72° north latitude in the Chukchi Sea; and on the south ...

n land bridge allowed members of the genus '' Eucyon'' a means to enter Asia from North America and they continued on to colonize Europe.

Pliocene epoch

The ''

Canis ''Canis'' is a genus of the Caninae which includes multiple extant taxon, extant species, such as Wolf, wolves, dogs, coyotes, and golden jackals. Species of this genus are distinguished by their moderate to large size, their massive, well-develo ...

'', '' Urocyon'', and ''

Vulpes '' Vulpes '' is a genus of the subfamily Caninae. The members of this genus are colloquially referred to as true foxes, meaning they form a proper clade. The word "fox" occurs in the common names of all species of the genus, but also appears in ...

'' genera developed from canids from

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

, where the canine radiation began. The success of these canids was related to the development of lower

s that were capable of both

and shearing. Around 5 million years ago, some of the Old World '' Eucyon'' evolved into the first members of ''Canis'', In the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Canis 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''. The formation of the

Isthmus of Panama The Isthmus of Panama, historically known as the Isthmus of Darien, is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North America, North and South America. The country of Panama is located on the i ...

, about 3 Mya, joined

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

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 The gray fox (''Urocyon cinereoargenteus''), or grey fox, is an omnivorous mammal of the family Canidae, widespread throughout North America and Central America. This species and its only congener (biology), congener, the diminutive island fox ...

(''Urocyon cinereoargentus'') and the now-extinct

dire wolf The dire wolf (''Aenocyon dirus'' ) is an Extinction, extinct species of Caninae, canine which was native to the Americas during the Late Pleistocene and Early Holocene epochs (125,000–10,000 years ago). The species was named in 1858, four y ...

(''Aenocyon dirus''). Besides these, there are species endemic to South America: the

maned wolf The maned wolf (''Chrysocyon brachyurus'') is a large canine of South America. It is found in Argentina, Brazil, Bolivia, Peru, and Paraguay, and is almost extinct in Uruguay. Its markings resemble those of a red fox, but it is neither a fox nor ...

(''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.

Pleistocene epoch

During the

Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

, the North American wolf line appeared, with '' Canis edwardii'', clearly identifiable as a wolf, and '' Canis rufus'' appeared, possibly a direct descendant of ''C. edwardii''. Around 0.8 Mya, '' 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. By 0.3 Mya, a number of subspecies of the gray wolf (''C. lupus'') had developed and had spread throughout Europe and northern Asia. 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. MtDNA studies have shown that there are at least four extant ''C. lupus'' lineages. 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. 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'' ( African golden wolf). According to a phylogeny derived from nuclear sequences, the Eurasian golden jackal (''Canis aureus'') diverged from the wolf/coyote lineage 1.9 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.

Wild canids are native to all continents except Australasia and Antarctica, and also occur as
feral A feral (; ) animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in som ...
(human-introduced) in New Guinea and Australia. They inhabit a wide range of different habitats, including
deserts A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the l ...
,
mountains A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher ...
,
forests A forest is an ecosystem characterized by a dense community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological functio ...
, and
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
s. They vary in size from the fennec fox, which may be as little as in length and weigh , to the gray
wolf The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a Canis, canine native to Eurasia and North America. More than thirty subspecies of Canis lupus, subspecies of ''Canis lupus'' have been recognized, includin ...
, which may be up to long, and can weigh up to .Heptner, V. G.; Naumov, N. P. (1998)
Mammals of the Soviet Union Vol. II Part 1a, Sirenia and Carnivora (Sea cows; Wolves and Bears)
Science Publishers, Inc. US., pp. 166–176, Only a few species are
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
—the
gray fox The gray fox (''Urocyon cinereoargenteus''), or grey fox, is an omnivorous mammal of the family Canidae, widespread throughout North America and Central America. This species and its only congener (biology), congener, the diminutive island fox ...
, the closely related island fox and the raccoon dog habitually climb trees. 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 dog breeds, canids have relatively long legs and lithe bodies, adapted for chasing prey. The tails are bushy and the length and quality of the pelage vary with the season. The muzzle portion of the skull is much more elongated than that of the cat family. The zygomatic arches are wide, there is a transverse lambdoidal ridge at the rear of the cranium and in some species, a
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are excepti ...
running from front to back. The bony orbits around the eye never form a complete ring and the auditory bullae are smooth and rounded. Females have three to seven pairs of mammae. 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 A dewclaw is a digit – vestigial in some animals – on the foot of many mammals, birds, and reptiles (including some extinct orders, like certain theropods). It commonly grows higher on the leg than the rest of the foot, such that in di ...
, 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 while other species have sharper, partially-retractile claws. The canine penis contains a
baculum The baculum (: bacula), also known as the penis bone, penile bone, ''os penis'', ''os genitale'', or ''os priapi'', is a bone in the penis of many placental mammals. It is not present in humans, but is present in the penises of some primates, ...
and a structure called the bulbus glandis that expands during copulation, forming a
copulatory tie Canine reproduction is the process of sexual reproduction in domestic dogs, wolves, coyotes and other canine species. Canine sexual anatomy and development Male reproductive system Erectile tissue As with all mammals, a dog's penis is made up ...
that lasts for up to an hour. Young canids are born blind, with their eyes opening a few weeks after birth. All living canids (Caninae) have a ligament analogous to the
nuchal ligament The nuchal ligament is a ligament at the back of the neck that is continuous with the supraspinous ligament. Structure The nuchal ligament extends from the external occipital protuberance on the skull and median nuchal line to the spinous p ...
of 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.

Dentition

Dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
relates to the arrangement of teeth in the mouth, with the dental notation for the upper-jaw teeth using the upper-case letters I to denote
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
, C for canines, P for premolars, and M for
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
, and the lower-case letters i, c, p and m to denote the mandible teeth. Teeth are numbered using one side of the mouth and from the front of the mouth to the back. In
carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
, the upper premolar P4 and the lower molar m1 form the carnassials that are used together in a scissor-like action to shear the muscle and tendon of prey. 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 trigonid is used for slicing and 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 A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant exampl ...
and a larger talonid indicating a more
omnivorous An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
diet. Because of its low variability, the length of the lower carnassial is used to provide an estimate of a carnivore's body size. 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. Most canids have 42
teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
, with a dental formula of: . The bush dog has only one upper molar with two below, the dhole has two above and two below. and the
bat-eared fox The bat-eared fox (''Otocyon megalotis'') is a species of fox found on the African savanna. It is the only extant species of the genus ''Otocyon'' and a Basal (phylogenetics), basal species of Canidae, canid. Fossil records indicate this canid ...
has three or four upper molars and four lower ones. 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 , molars being completely absent.

Life history

Social behavior
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. 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. 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.

Communication
Canids communicate with each other by scent signals, by visual clues and gestures, and by vocalizations such as growls, barks, and howls. In most cases, groups have a home territory from which they drive out other
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
s. Canids use urine scent marks to mark their food caches or warn trespassing individuals. Social behavior is also mediated by secretions from glands on the upper surface of the tail near its root and from the anal glands, preputial glands, and supracaudal glands.

Reproduction
Canids as a group exhibit several reproductive traits that are uncommon among mammals as a whole. They are typically monogamous, provide paternal care to their offspring, have reproductive cycles with lengthy proestral and dioestral phases and have a copulatory tie during mating. They also retain adult offspring in the social group, suppressing the ability of these to breed while making use of the alloparental care they can provide to help raise the next generation. Most canid species are spontaneous ovulators, though
maned wolves The maned wolf (''Chrysocyon brachyurus'') is a large caninae, canine of South America. It is found in Argentina, Brazil, Bolivia, Peru, and Paraguay, and is almost extinct in Uruguay. Its markings resemble those of a red fox, but it is neither a ...
are induced ovulators. During the proestral period, increased levels of estradiol make the female attractive to the male. There is a rise in
progesterone Progesterone (; P4) is an endogenous steroid and progestogen sex hormone involved in the menstrual cycle, pregnancy, and embryogenesis of humans and other species. It belongs to a group of steroid hormones called the progestogens and is the ma ...
during the estral phase when female is receptive. Following this, the level of estradiol fluctuates and there is a lengthy dioestrous phase during which the female is pregnant. Pseudo-pregnancy often occurs in canids that have ovulated but failed to conceive. A period of anestrus follows pregnancy or pseudo-pregnancy, there being only one oestral period during each breeding season. Small and medium-sized canids mostly have a
gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregn ...
of 50 to 60 days, while larger species average 60 to 65 days. The time of year in which the breeding season occurs is related to the length of day, as has been shown for several species that have been moved across the equator and experiences a six-month shift of phase. Domestic dogs and certain small canids in captivity may come into oestrus more often, perhaps because the photoperiod stimulus breaks down under conditions of artificial lighting. Canids have an oestrus period of 1 to 20 days, lasting one week in most species. 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. 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. Young canids may take a year to mature and learn the skills they need to survive. 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.

Canids and humans
One canid, the
domestic dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers ...
, entered into a partnership with humans a long time ago. The dog was the first domesticated species. The archaeological record shows the first undisputed dog remains buried beside humans 14,700 years ago, with disputed remains occurring 36,000 years ago. These dates imply that the earliest dogs arose in the time of human
hunter-gatherer A hunter-gatherer or forager is a human living in a community, or according to an ancestrally derived Lifestyle, lifestyle, in which most or all food is obtained by foraging, that is, by gathering food from local naturally occurring sources, esp ...
s and not agriculturists. 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. Among canids, only the gray wolf has widely been known to prey on humans. Nonetheless, at least two records of coyotes killing humans have been published, and at least two other reports of golden jackals killing children. 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. Canids such as the dhole are now
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
in the wild because of persecution, habitat loss, a depletion of ungulate prey species and transmission of diseases from domestic dogs.

See also
* List of canids * Largest wild canids *
Canid hybrid Canid hybrids are the result of interbreeding between the species of the subfamily ''Caninae''. Genetic considerations The wolf-like canids are a group of large carnivores that are genetically closely related because they all possess 78 chromos ...
* Free-ranging dog

References

Bibliography
*

External links
* {{Authority control Mammal families Bartonian first appearances Extant Eocene first appearances Taxa named by Gotthelf Fischer von Waldheim