Canidae

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. The Canidae first appeared in North America during the Late Eocene (37.8-33.9 Mya). They did not reach Eurasia until the Late Miocene or to South America until the Late Pliocene. ==Evolution==

(Hesperocyoninae), Aelurodon (Borophaginae) and 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. Eocene epoch Carnivorans evolved after the extinction of the non-avian dinosaurs 66 million years ago. Around 50 million years ago, or earlier, in the Paleocene, the Carnivora split into two main divisions: caniform (dog-like) and 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. 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 Caninae (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, 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. 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. Pliocene epoch The Canis, Urocyon, and Vulpes genera developed from canids from North America, where the canine radiation began. The success of these canids was related to the development of lower carnassials that were capable of both mastication and shearing. 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. 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. 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. ====

{{multiple image|direction=vertical|image1=|caption1=Comparative illustration of the paws of the wolf, golden jackal, and dhole by A. N. Komarov| Wild canids are native to all continents except Australasia and Antarctica, and also occur as feral (human-introduced) in New Guinea and Australia. They inhabit a wide range of different habitats, including deserts, mountains, forests, and grasslands. They vary in size from the fennec fox, which may be as little as in length and weigh , to the gray wolf, which may be up to long, and can weigh up to . Only a few species are arboreal—the gray 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 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, 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 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 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 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, C for canines, P for premolars, and M for molars, 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, 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 and a larger talonid indicating a more omnivorous 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, 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 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 {{multiple image|perrow=1|image1=2012-bandipur-dhole-sambar.jpg|caption1=Dholes attacking a sambar, Bandipur National Park| 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 conspecifics. 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, 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 are induced ovulators. During the proestral period, increased levels of estradiol make the female attractive to the male. There is a rise in progesterone 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 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. 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. 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, entered into a partnership with humans a long time ago. The dog was the first domesticated species. 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 in the wild because of persecution, habitat loss, a depletion of ungulate prey species and transmission of diseases from domestic dogs. ==See also==