Canidae

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Canids[1]
Temporal range: 39.75–0Ma
Late Eocene - Recent
Major extant canid genera: Canis, Cuon, Lycaon, Cerdocyon, Chrysocyon, Speothos, Vulpes, Nyctereutes, Otocyon and Urocyon
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Family: Canidae
G. Fischer de Waldheim, 1817
Genera and species

See text

The Canidae (pron.: /ˈkænɨdiː/) [2] are the biological family of carnivorous and omnivorous mammals that includes domestic dogs, wolves, foxes, jackals, coyotes, and many other lesser known extant and extinct dog-like mammals. A member of this family is called a canid (/ˈkeɪnɨd/). The Canidae family is divided into two tribes: Canini (related to wolves) and Vulpini (related to foxes). The two species of the basal Caninae are more primitive and do not fit into either tribe.

Contents

Classification and relationship [edit]

The subdivision of Canidae into "foxes" and "true dogs" may not be in accordance with the actual relations; also, the taxonomic classification of several canines is disputed. Recent DNA analysis shows that Canini (dogs) and Vulpini (foxes) are valid clades (see phylogeny below). Molecular data imply a North American origin of living Canidae and an African origin of wolf-like canines (Canis, Cuon, and Lycaon).[3]

Currently, the domestic dog is listed as a subspecies of Canis lupus, C. l. familiaris, and the dingo (also considered a domestic dog) as C. l. dingo, provisionally a separate subspecies from C. l. familiaris; the red wolf, eastern Canadian wolf, and Indian wolf are recognized as subspecies.[1] Many sources list the domestic dog as Canis familiaris, but others, including the Smithsonian Institution and the American Society of Mammalogists, more precisely list it as a subspecies of C. l. familiaris; the red wolf, eastern Canadian wolf, and Indian wolf may or may not be separate species; in the past, the dingo has been variously classified as Canis dingo, Canis familiaris dingo and Canis lupus familiaris dingo.

Evolution [edit]

Evolution of the Canids
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Axis scale: millions of years ago.
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Eocene epoch [edit]

Carnivorans evolved from miacoids about 55 million years ago (Mya) during the late Paleocene.[4] Then, about 50 Mya, the carnivorans split into two main divisions: caniforms (dog-like) and feliforms (cat-like). By 40 Mya, the first clearly identifiable member of the dog family, Canidae, had arisen. Called Prohesperocyon wilsoni, it was found in what is now southwestern Texas. This fossil species bears a combination of features that definitively mark it as a canid: teeth that include the loss of the upper third molar (a general trend toward a more shearing bite), and the characteristically enlarged bony bulla (the rounded covering over the middle ear). Based on what is known about its descendants, Prohesperocyon likely had slightly more elongated limbs than its predecessors, along with parallel and closely touching toes, rather than splayed as in bears.[5]

The Canidae family soon subdivided into three subfamilies, each of which diverged during the Eocene: Hesperocyoninae (about 39.74-15 Mya), Borophaginae (about 34-2 Mya), and the Caninae (about 34-0 Mya) lineage that led to present-day canids (wolves, foxes, coyotes, jackals, and domestic dogs). Each of these groups showed an increase in body mass with time, and sometimes exhibited specialised hypercarnivorous diets that made them prone to extinction.[6]:Fig. 1 Only the Caninae lineage, commonly referred to as "canines", survived to the present day.

Oligocene epoch [edit]

By the Oligocene, all three subfamilies of canids (Hesperocyoninae, Borophaginae, and Caninae) had appeared in the fossil records of North America. The earliest and most primitive branch of the Canidae was the Hesperocyoninae lineage, 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 civets in appearance. Hesperocyonines eventually became extinct in the middle Miocene. One of the early member of the Hesperocyonines, the genus Hesperocyon, gave rise to Archaeocyon and Leptocyon. These branches led to the borophagine and canine radiations.[7]

Miocene epoch [edit]

Around 9-10 Mya during the Late Miocene, Canis, Urocyon, and Vulpes genera expanded from southwestern North America, where the canine radiation began. The success of these canines was related to the development of lower carnassials that were capable of both mastication and shearing. Around 8 Mya, Beringia offered the canines a way to enter Eurasia.

Pliocene epoch [edit]

During the Pliocene, around 4-5 Mya, Canis lepophagus appeared in North America. This was small and sometimes coyote-like. Others were wolf-like in characteristics.Canis latrans (the coyote) is theorized to have descended from Canis lepophagus.[8]

The formation of the Isthmus of Panama, about 3 Mya, joined South America to North America, allowing canids to invade the former, where they diversified.

Pleistocene epoch [edit]

Around 1.5 to 1.8 Mya, a variety of wolves were present in Europe.[citation needed] Also, the North American wolf line appeared with Canis edwardii, clearly identifiable as a wolf. Canis rufus appeared, possibly a direct descendent of Canis 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 its descendant, Canis dirus, the dire wolf, which then also spread into South America during the late Pleistocene.

By 0.3 Mya, Canis lupus was fully developed and had spread throughout Europe and northern Asia, and Beringia offered a way to North America.[9] At around 100,000 years ago, the dire wolf, one of the largest members of the dog family, had spread from South America to southern Canada and from coast to coast. The dire wolf shared its habitat with the gray wolf. Around 8,000 years ago, the dire wolf became extinct.

Characteristics [edit]

Wild canids are found on every continent except Antarctica, and inhabit a wide range of different habitats, including deserts, mountains, forests, and grasslands. They vary in size from the fennec fox at 24 cm (9.4 in) long, to the gray wolf, which may be up to 2 m (6.6 ft) long, and can weigh up to 80 kg (180 lb).

With the exceptions of the bush dog, raccoon dog, and some domestic breeds of Canis lupus, canids have relatively long legs and lithe bodies, adapted for chasing prey. All canids are digitigrade, meaning they walk on their toes. They possess bushy tails, nonretractile claws, and, excepting the African wild dog (Lycaon pictus), a dewclaw on the front feet. They possess a baculum that, together with a cavernous body, helps to create a copulatory tie during mating, locking the animals together for up to an hour. Young canids are born blind, with their eyes opening a few weeks after birth.[10] 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.[11] However, based on skeletal details of the neck, at least some Borophaginae (such as Aelurodon) are believed to have lacked this ligament.[11]

Gray wolf pack hunting an American bison in Yellowstone National Park.

Only a few species are arboreal - the North American gray fox, the closely related Channel Island fox,[12] and the raccoon dog habitually climb trees.[13][14][15]

Social behavior [edit]

See also: Gray wolf#Behaviour, Dog behavior, Red fox#Behaviour, and African Wild Dog#Social structure

Almost all canids are social animals and live together in groups. 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 the largest packs, which can number as many as 90 animals. 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.

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 others. The territory is marked by leaving urine scent marks, which warn trespassing individuals.[16]

Territorial marking [edit]

Wikimedia Commons has media related to: Urinating canids
See also: Canidae#Social behavior
A dog marking his territory

All canids (with the possible exception of dholes[17]) use urine (combined with preputial gland secretions) to mark their territories. Many species of canids, including hoary foxes,[18] cape foxes,[19] and golden jackals,[20] use a raised-leg posture when urinating.[21][22] The scent of their urine is usually strongest in the winter, before the mating season.[22]

Domestic dogs mark their territories by urinating on vertical surfaces (usually at nose level), sometimes marking over the urine of other dogs.[23] When one dog marks over another dog's urine, this is known as "counter-marking" or "overmarking".[24][25] Male dogs urine-mark more frequently than female dogs,[26] typically beginning after the onset of sexual maturity.[27] Male dogs, as well as wolves, sometimes lift a leg and attempt to urinate even when their bladders are empty - this is known as a "raised-leg display"[28] or "pseudo-urination".[29][30] They typically mark their territory due to the presence of new stimuli or social triggers in a dog's environment, as well as out of anxiety.[31] Marking behavior is present in both male dogs and female dogs, and are especially pronounced in male dogs that have not been neutered.[31]

Raised-leg urination is the most significant form of scent marking in wolves, and is most frequent around the breeding season.[32] Wolves urine-mark more frequently when they detect the scent of other wolves, or other canid species.[33] Leg-lifting is more common in male wolves than female wolves, although dominant females also use the raised-leg posture.[34]

A wolf marking its territory

Other types of urine-marking in wolves are FLU (flexed-leg urination), STU (standing urination), and SQU (squatting urination).[35] Breeding pairs of wolves will sometimes urinate on the same spot: this is known as "double-marking".[36][37][38][39][40][41] Double-marking is practiced by coyotes as well as wolves.,[42][43][44] and also by foxes.[45]

Coyotes mark their territories by urinating on bushes, trees, or rocks.[46] All male coyotes lift their legs when urinating.[47] However, females sometimes also raise their legs, and males sometimes squat.[48] Urine marking is also associated with pair bonding in coyotes[clarification needed][49] Coyotes sometimes urinate on their food, possibly to claim ownership over it.[50]

Red foxes use their urine to mark their territories.[51][52][53][54][55] A male fox raises one hind leg and his urine is sprayed forward in front of him, whereas a female fox squats down so that the urine is sprayed in the ground between the hind legs.[56][57] Urine is also used to mark empty cache sites, as reminders not to waste time investigating them.[58][59][60] Red foxes use various postures[clarify] to urinate, depending on where they are leaving a scent mark.[56][61]

As in most other canids, male bush dogs lift their hind legs when urinating. However, female bush dogs use a kind of handstand posture, which is less common in other canids.[62][63] When male bush dogs urinate, they create a spray instead of a stream.[64]

Both male and female maned wolves use their urine to communicate, e.g. to mark their hunting paths, or the places where they have buried hunted prey.[65] The urine has a very distinctive smell, which some people liken to hops or cannabis. The responsible substance is very likely a pyrazine, which occurs in both plants.[66] (At the Rotterdam Zoo, this smell once set the police on a hunt for cannabis smokers.[66][67])


Reproduction [edit]

[icon] This section requires expansion. (January 2013)
See also: Canine reproduction and Animal sexual behavior#Canidae

Most canids bear young once a year, from one to 16 or more (in the case of the African wild dog) at a time. The young are born small and helpless and require a long period of care. They are kept in a den, most often dug into the ground, for warmth and protection. When they 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. Young canids may take a year to mature and learn the skills they need to survive.[68]

Reproductive anatomy [edit]
[icon] This section requires expansion. (January 2013)

In male canids, the penis is highly specialized[clarification needed], and is supported by a bone called the baculum. It also contains a structure at the base called the bulbus glandis.[69][further explanation needed]

Dentition [edit]

Eurasian wolf skull

Most canids have 42 teeth, with a dental formula of: 3.1.4.23.1.4.3. As in other members of Carnivora, the upper fourth premolar and lower first molar are adapted as carnassial teeth for slicing flesh. 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 3; molars are completely absent.

Species and taxonomy [edit]

The genus Canis: Grey wolf, coyote, golden jackal, and Ethiopian wolf
The genus Lycalopex: Culpeo, pampas fox, chilla and Darwin's fox
The genus Vulpes: red fox, Rüppell's fox, corsac fox, Bengal fox, Arctic fox, Blanford's fox, Cape fox and fennec fox
Dhole
African hunting dog
Short-eared dog
Maned wolf
Bush dog
Gray fox
Bat-eared fox
Raccoon dog

FAMILY CANIDAE (Extant and recently extinct species)

Subfamily Caninae [edit]

Fluctuation of species within Canidae over 40 million years

Prehistoric Canidae [edit]

Classification of Hesperocyoninae from Wang (1994);[70] Borophaginae from Wang, Tedford, Taylor (1999),;[71] and Caninae from Tedford, Wang, Taylor (2009)[72] except where noted.

Caninae [edit]

Borophaginae [edit]

† (Mya = million years ago) (million years = in existence)

Hesperocyoninae [edit]

† (Mya = million years ago)

    • Genus Cynodesmus (32-29 Mya)
      • Cynodesmus martini (29 Mya)
      • Cynodesmus thooides (32 Mya)
    • ?Genus Caedocyon
      • Caedocyon tedfordi
    • Genus Ectopocynus (32-19 Mya)
      • Ectopocynus antiquus (32 Ma)
      • Ectopocynus intermedius (29 Mya)
      • Ectopocynus siplicidens (19 Mya)
    • Genus Enhydrocyon (29-25 Mya)
      • Enhydrocyon basilatus (25 Mya)
      • Enhydrocyon crassidens (25 Mya)
      • Enhydrocyon pahinsintewkpa (29 Mya)
      • Enhydrocyon stenocephalus (29 Mya)
    • Genus Hesperocyon (39.74-34 Mya)
      • Hesperocyon coloradensis
      • Hesperocyon gregarius (37 Mya)
    • Genus Mesocyon (34-29 Mya)
      • Mesocyon brachyops (29 Mya)
      • Mesocyon coryphaeus (29 Mya)
      • Mesocyn temnodon
    • Genus Osbornodon (32-18 Mya)
      • Osbornodon brachypus
      • Osbornodon fricki (18 Mya)
      • Osbornodon iamonensis (21 Mya)
      • Osbornodon renjiei (33 Mya)
      • Osbornodon scitulus[75]
      • Osbornodon sesnoni (32 Mya)
      • Osbornodon wangi[73]
    • Genus Paraenhydrocyon (30-25 Mya)
    • Genus Philotrox (29 Mya)
      • Philotrox condoni (29 Mya)
    • Genus Prohesperocyon (36 Mya)
      • Prohesperocyon wilsoni (36 Mya)
    • Genus Sunkahetanka (29 Mya)
      • Sunkahetanka geringensis (29 Mya)

Canids and humans [edit]

Traditional English fox hunt

One canid, the domestic dog, a subspecies of the gray wolf, long ago entered into a partnership with humans, and today remains one of the most widely kept domestic animals in the world and serves humanity in many important ways. This partnership is documented as far back as 26,000 years ago, when the footprints of a young boy about the age of eight to 10 years old was found in Chauvet Cave in southern France, alongside of what was identified as a large dog or wolf.[76]

The earliest recorded fossil of a dog was found around 31,700 years ago in Goyet Cave in Belgium.[77] This is indicative of the relationship canids and humans had during the evolution of wolves to dogs. Still even early on, wolves were found fossilized with humans that date back 300,000 years ago, which shows how far back humans and wolves had interaction with one another.[78] The bond most pet owners have towards their dogs can be traced back early on in the dog. This affection can be seen with the burial of dogs with humans as early as 11,000 years ago in the Americas and 8,500 years ago in Europe.[78]

Among canids, only the gray wolf has widely been known to prey on humans.[79] Nonetheless, at least two records have coyotes killing humans,[80] and two have golden jackals killing children.[81] Human beings have trapped and hunted some canid species for their fur and, especially the gray wolf and the red fox, for sport. Some canids are now endangered in the wild because of hunting, habitat loss, and introduction of diseases from domestic dogs.[82]

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General references [edit]

  • Xiaoming Wang, Richard H. Tedford, Mauricio Antón, Dogs: Their Fossil Relatives and Evolutionary History, New York : Columbia University Press, 2008; ISBN 978-0-231-13528-3

External links [edit]

Wikispecies has information related to: Canidae
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