Canidae

Taxobox
name = CanidsMSW3 Wozencraft | pages = | id = 14000691]
fossil_range = Paleogene (55.4 Ma) - Recent

The Canidae (IPAEng|ˈkænədiː, ′kanə′dē) family is a part of the order Carnivora within the mammals (Class Mammalia). Members of the family are called canids and include dogs, wolves, foxes, coyotes, dingoes, jackals, and African Wild Dogs. The Canidae family is divided into the "true dogs" (or canines) of the tribe Canini and the "foxes" of the tribe Vulpini. The two species of the basal Caninae are more primitive and don't fit into either tribe. Any member of this family can be referred to generally as a canid.

Classification and relationship

Note that the subdivision of Canidae into "foxes" and "true dogs" may not be in accordance with the actual relations, and that the taxonomic classification of several canines is disputed. Recent DNA analysis has shown, however, that Canini (dogs) and Vulpini (foxes) are valid clades. (See phylogeny below). Molecular data implies a North American origin of living Canidae and an African origin of wolf-like canines ("Canis", "Cuon", and "Lycaon").cite journal | author = Lindblad-toh, K. | coauthors = Wade, C.M.; Mikkelsen, T.S.; Karlsson, E.K.; Jaffe, D.B.; Kamal, M.; Clamp, M.; Chang, J.L.; Kulbokas Iii, E.J.; Zody, M.C.; Others, | year = 2005 | title = Genome sequence, comparative analysis and haplotype structure of the domestic dog | journal = Nature | volume = 438 | issue = 7069 | pages = 803–819 | url = http://ccr.cancer.gov/resources/cop/nature04338.pdf | accessdate = 2008-04-27 | doi = 10.1038/nature04338]

Currently the domestic dog is listed as a subspecies of "Canis lupus", "C. l. familiaris", with the Dingo (also considered a domestic dog) listed 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 as well.

The domestic dog is listed by some authoritiesWho|date=May 2008 as "Canis familiaris" and others (including the Smithsonian InstitutionFact|date=May 2008 and the American Society of MammalogistsFact|date=May 2008) as a subspecies of the Gray Wolf (i.e., "Canis lupus familiaris"); the Red Wolf, Eastern Canadian Wolf, and Indian Wolf may or may not be separate species; and the Dingo has been in the past variously classified as "Canis dingo", "Canis familiaris dingo" and "Canis lupus familiaris dingo".

Evolution

Eocene epoch

The Canidae family evolved from miacids about 40 million years ago in the late Eocene to early Oligocene. The Canidae family is subdivided into three subfamilies, each of which diverged during the Eocene: Hesperocyoninae (~39.74-15 Mya), Borophaginae (~36-2 Mya), and the Caninae lineage that led to present-day canids, including wolves, foxes, coyotes, jackals, and domestic dogs.

Each of the groups showed an increase in body mass with time before their specialised hypercarnivorous diet made them prone to extinction. [cite doi|10.1126/science.1102417] rp|Fig. 1

Oligocene epoch

The earliest 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 fast pursuit of prey in a grassland habitat, and resembled modern civets in appearance. Hesperocyonine dogs became extinct except for the "Nothocyon" and "Leptocyon" branches. These branches lead 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 9-10 Mya during the Late Miocene, "Canis", "Urocyon", and "Vulpes" genera expand from southwestern North America. This is the point where canine radiation begins. The success of the these canines is the development of lower carnassials that are capable of both mastication and shearing. Around 8 Mya, Berengia offers the canines a way to enter Eurasia.

Pliocene epoch

Early Pliocene

During the Pliocene around (4-5 Mya) "Canis lepophagus" appears in North America. This dog is small with some being coyote-like. Others are wolf-like in characteristics. It is theorized that "Canis latrans" (coyote) descended from "Canis lepophagus". [Nowak, R.M. 1979. North American Quaternary Canis. Monograph of the Museum of Natural History, University of Kansas 6:1 - 154.] Around 1.5 to 1.8 Mya, a variety of wolves are now in Europe. Also, the North American wolf line appears with "Canis edwardii" as clearly identifiable as a wolf. "Canis rufus", a red wolf canine appears and possibly a direct descendent of Canis edwardii.

Middle Pliocene

Around 0.8 Mya "Canis ambrusteri", emerges in North America. A large wolf, it is found all over the continent. It is thought that this species went to South America where it becomes the ancestor of the "Canis dirus" or Dire wolf.

Late Pliocene

At 0.3 Mya "Canis lupus" (Gray wolf) has fully developed and has spread throughout Europe and northern Asia. Berengia offers a way to North America. [Nowak, R. 1992. Wolves: The great travelers of evolution. International Wolf 2(4):3 - 7.] At around 100,000 years ago, the Dire wolf, some of the largest members of the dog family, appears from southern Canada to South America and coast to coast. The Dire wolf shares its habitat with the Gray wolf. Around 8000 years ago the Dire wolf becomes extinct.

Characteristics

Wild canids are found on every continent except Antarctica, and inhabit a wide range of different habitats, including deserts, mountains, forests, and grassland. They vary in size from the Fennec Fox at 24 cm in length, to the Gray Wolf, which may be up to 200 cm long, and can weigh up to 80 kg.

With the sole living exception of the Bush Dog, canids have relatively long legs and lithe bodies, adapted for chasing prey. All canids are digitigrade, meaning that they walk on their toes. They possess bushy tails, non-retractile claws, and a dewclaw on the front feet. They possess a baculum, which 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. [cite book |author=Macdonald, D.|year=1984 |title= The Encyclopedia of Mammals|publisher= Facts on File|location=New York|pages= 57|isbn= 0-87196-871-1]

Many species live and hunt in packs, and have complex social lives. They are generally highly adaptable, and there may be considerable variation in habits even within a single species.

Dentition

Most canids have 42 teeth, with a dental formula of: dentition2|3.1.4.2|3.1.4.3 As in other members of the 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.

pecies and taxonomy

FAMILY CANIDAE

Subfamily: Caninae
* True dogs - Tribe Canini
** "Genus Canis"
*** Side-striped Jackal, "Canis adustus"
*** Golden Jackal, "Canis aureus"
*** Coyote, "Canis latrans" (also called Prairie Wolf)
*** Gray Wolf, "Canis lupus" (2.723 Ma to present)
**** Red Wolf, "Canis lupus rufus" (3 Ma to present) (sometimes considered a separate species)
**** Domestic Dog, "Canis lupus familiaris"
**** Dingo, most often classified as "Canis lupus dingo"
**** New Guinea Singing Dog, "Canis lupus hallstromi"
**** Eastern Wolf, "Canis (lupus) lycaon" (sometimes considered a separate species)
**** Indian Wolf, "Canis indica" (sometimes considered a separate species)
**** Himalayan Wolf, "Canis himalayensis" (sometimes considered a separate species)
**** many other proposed subspecies
*** Black-backed Jackal, "Canis mesomelas"
*** Ethiopian Wolf, "Canis simensis" (also called Abyssinian Wolf, Simien Fox and Simien Jackal)
** "Genus Cynotherium" †
*** Sardinian Dhole, "Cynotherium sardous" †
** "Genus Cuon"
*** Dhole, "Cuon alpinus" or "Canis alpinus" (also called Asian Wild Dog)
** "Genus Lycaon"
*** African Wild Dog, "Lycaon pictus" (also called African Hunting Dog)
** "Genus Indocyon"†
*** Indian Mute Dog, "Indocyon caribensis" † (also called Caribbean Dog)
** "Genus Cubacyon"
*** Cuban Dhole, "Cubacyon transversidens" †
** "Genus Atelocynus"
*** Short-eared Dog, "Atelocynus microtis"
** "Genus Cerdocyon"
*** Crab-eating Fox, "Cerdocyon thous"
** "Genus Dasycyon" † ?
*** Hagenbeck Wolf, "Dasycyon hagenbecki" † ?
** "Genus Dusicyon" †
*** Falkland Island Fox, "Dusicyon australis" †
** "Genus Pseudalopex"
*** Culpeo, "Pseudalopex culpaeus"
*** Darwin's Fox, "Pseudalopex fulvipes"
*** Argentine Grey Fox, "Pseudalopex griseus"
*** Pampas Fox, "Pseudalopex gymnocercus"
*** Sechura Fox, "Pseudalopex sechurae"
*** Hoary Fox, "Pseudalopex vetulus"
** "Genus Chrysocyon"
*** Maned Wolf, "Chrysocyon brachyurus"
** "Genus Speothos"
*** Bush Dog, "Speothos venaticus"
*** unnamed bush dog species, "Speothos pacivorus" †

* True foxes - Tribe Vulpini
** "Genus Vulpes"
*** Arctic Fox, "Vulpes lagopus"
*** Red Fox, "Vulpes vulpes" (1 Ma to present)
*** Swift Fox, "Vulpes velox"
*** Kit Fox, "Vulpes macrotis"
*** Corsac Fox, "Vulpes corsac"
*** Cape Fox, "Vulpes chama"
*** Pale Fox, "Vulpes pallida"
*** Bengal Fox, "Vulpes bengalensis"
*** Tibetan Sand Fox, "Vulpes ferrilata"
*** Blanford's Fox, "Vulpes cana"
*** Rüppell's Fox, "Vulpes rueppelli"
*** Fennec Fox, "Vulpes zerda"
** "Genus Urocyon" (2 Ma to present)
*** Gray Fox, "U. cinereoargenteus"
*** Island Fox, "U. littoralis"
*** Cozumel Fox, "U". sp.

* Basal Caninae
** "Genus Otocyon" (probably a vulpine close to "Urocyon")
*** Bat-eared Fox, "Otocyon megalotis"
** "Genus Nyctereutes"
*** Raccoon Dog, "Nyctereutes procyonoides"

Fossil Canidae

Classification of Hesperocyoninae from Wang (1994)cite journal| last = Wang| first = Xiaoming| year = 1994 |title = Phylogenetic systematics of the Hesperocyoninae| journal = Bulletin of the American Museum of Natural History| volume = 221| pages= 1–207| url = http://hdl.handle.net/2246/829] and Borophaginae from Wang et al. (1999),cite journal| last = Wang| first = Xiaoming| coauthors = Richard Tedford, Beryl Taylor| year = 1999 | title = Phylogenetic systematics of the Borophaginae| journal = Bulletin of the American Museum of Natural History| volume = 243| pages= 1–391| url = http://hdl.handle.net/2246/1588 | doi = 10.1671/2493] except where noted.

Prehistoric Caninae
*Canini
** "Genus Canis"
*** Dire Wolf, "Canis dirus" (1 Ma †)
*** "Canis arnensis" (3.4 Ma, †)
*** "Canis (Eucyon) cipio" (8.2 Ma †, probably first species of "Canis" genus)
*** "Canis etruscus" (3.4 Ma †)
*** "Canis mosbachensis" (0.787 Ma †)
*** "Canis lepophagus" (4-5 Ma †)
*** "Canis donnezani" (4.0-3.1 Ma †, probably ancestor of wolves)
*** "Canis edwardii" (1.8 Ma †, first species of wolf in North America)
*** "Canis gezi" †
*** "Canis nehringi" †
*** "Canis ameghinoi " †
*** "Canis michauxi " †
*** "Canis adoxus " †
*** "Canis cautleyi " †
*** "Canis armbrusteri " (0.8 Ma †)
** "Genus Theriodictis" (1.8 Ma †)
*** "Theriodictis platensis" (1.8 Ma †)
*** "Theriodictis tarijensis" (1.8 Ma †)
*** "Theriodictis (Canis) proplatensis" (2.1 Ma † )
** "Genus Protocyon" †
*** "Protocyon orcesi" †
*** "Protocyon scagliarum" †
*** "Protocyon troglodytes" †
** "Genus Dusicyon" †
*** "Dusicyon avus" †
** "Genus Cerdocyon"
*** "Cerdocyon avius" †
*** "Cerdocyon ensenadensis" †
** "Genus Speothos"
*** "Speothos pacivorus" †
** "Genus Nurocyon"
*** "Nurocyon chonokhariensis" †
** "Genus Xenocyon"
*** "Xenocyon falconeri" (2.6 Ma †)
*** "Xenocyon lycaonoides"

*Vulpini
** "Genus Vulpes" (7 Ma to present)
*** "Vulpes alopecoides" (2.6 Ma †)
*** "Vulpes cf. alopecoides" (2.6 Ma †)
*** "Vulpes cf. vulpes" (0.1275 Ma †)
*** "Vulpes galaticus" (4.2 Ma †)
*** "Vulpes riffautae" (7 Ma †)
*Basal Canids
** "Genus Nyctereutes" (7.1 Ma to present)
*** "Nyctereutes cf. donnezani" (7.1 Ma †)
*** "Nyctereutes cf. megamastoides" (3.158 Ma †)
*** "Nyctereutes donnezani" (3.4 Ma †)
*** "Nyctereutes megamostoides" (2.6 Ma †)
*** "Nyctereutes sinensis" (3.4 Ma †)

*First Caninae
** "Genus Eucyon" (8 Ma †)
*** "Eucyon davisi" (8.3 Ma †, probably ancestor of "Canis")
*** "Eucyon minor" (8 Ma †)
*** "Eucyon zhoui" (8 Ma †)
*** "Eucyon monticinensis"(8 Ma †)
*** "Eucyon odessanus" †
** "Genus Leptocyon" (24-16 Ma †)
*** "Leptocyon vafer" (16 Ma)
*** "Leptocyon vulpinus" (24 Ma)

Borophaginae : † (Ma = million years ago)
** "Genus Aelurodon" (16-12 Ma)
*** "Aelurodon asthenostylus" (16 Ma)
*** "Aelurodon ferox" (15 Ma)
*** "Aelurodon mcgrewi" (15 Ma)
*** "Aelurodon montanensis" (15 Ma)cite journal| last = Wang| first = Xiaoming| coauthors = Wideman, Benjamin, Nichols, Ralph, & Hanneman, Debra| year = 2004| title = A new species of "Aelurodon" (Carnivora, Canidae) from the Barstovian of Montana| journal = Journal of Vertebrate Paleontology| volume = 24| issue = 2| pages = 445–452| url = http://www.nhm.org/expeditions/rrc/wang/documents/Wangetal2004MontanaAelurodon.pdf| accessdate = 2007-07-08| doi = 10.1671/2493 ]
*** "Aelurodon stirtoni" (13 Ma)
*** "Aelurodon taxoides" (12 Ma)
** "Genus Archaeocyon" (32-24 Ma)
*** "Archaeocyon falkenbachi" (25-24 Ma)
*** "Archaeocyon leptodus" (32-24 Ma)
*** "Archaeocyon pavidus" (32-28 Ma)
** "Genus Borophagus" (12-5 Ma)
*** "Borophagus dividersidens" (5 Ma)
*** "Borophagus dudleyi"
*** "Borophagus hilli" (6 Ma)
*** "Borophagus littoralis" (12 Ma)
*** "Borophagus orc" (9 Ma)
*** "Borophagus parvus" (7 Ma)
*** "Borophagus pugnator" (9 Ma)
*** "Borophagus secundus" (9 Ma)
** "Genus Carpocyon"
*** "Carpocyon compressus"
*** "Carpocyon limosus"
*** "Carpocyon robustus"
*** "Carpocyon webbi"
** "Genus Cormocyon"
*** "Cormocyon copei"
*** "Cormocyon haydeni"
** "Genus Cynarctoides" (30-18 Ma)
*** "Cynarctoides acridens" (24 Ma)
*** "Cynarctoides emryi" (21 Ma)
*** "Cynarctoides gawnae" (18 Ma)
*** "Cynarctoides harlowi" (21 Ma)
*** "Cynarctoides lemur" (30 Ma)
*** "Cynarctoides luskensis" (21 Ma)
*** "Cynarctoides roii" (30 Ma)
** "Genus Cynarctus" (16-12 Ma)
*** "Cynarctus crucidens" (12 Ma)
*** "Cynarctus galushai" (16 Ma)
*** "?Cynarctus marylandica"
*** "Cynarctus saxatilis" (15 Ma)
*** "Cynarctus voorhiesi" (13 Ma)
** "Genus Desmocyon" (24-19 Ma)
*** "Desmocyon matthewi" (19 Ma)
*** "Desmocyon thompsoni" (24 Ma)
** "Genus Epicyon" (12-10 Ma)
*** "Epicyon aelurodontoides" (10.3-4.9 Ma)
*** "Epicyon haydeni" (10 Ma)
*** "Epicyon saevus" (12 Ma)
** "Genus Eulopocyon" (18-16 Ma)
*** "Eulopocyon brachygnathus" (16 Ma)
*** "Eulopocyon spissidens" (18 Ma)
** "Genus Metatomarctus" (19-16 Ma)
*** "Metatomarctus canavus" (19 Ma)
*** "Metatomarctus sp. A" (16 Ma)
*** "Metatomarctus sp. B" (16 Ma)
** "Genus Microtomarctus" (18 Ma)
*** "Microtomarctus conferta" (18 Ma)
** "Genus Otarocyon" (34-30 Ma)
*** "Otarocyon cooki" (30 Ma)
*** "Otarocyon macdonaldi" (34 Ma)
** "Genus Oxetocyon" (32 Ma)
*** "Oxetocyon cuspidatus" (32 Ma)
** "Genus Paracynarctus" (19-16 Ma)
*** "Paracynarctus kelloggi" (19 Ma)
*** "Paracynarctus sinclairi" (16 Ma)
** "Genus Paratomarctus" (16-13 Ma)
*** "Paratomarctus euthos" (13 Ma)
*** "Paratomarctus temerarius" (16 Ma)
** "Genus Phlaocyon" (30-19 Ma)
*** "Phlaocyon achoros"
*** "Phlaocyon annectens" (22 Ma)
*** "Phlaocyon latidens" (30 Ma)
*** "Phlaocyon leucosteus" (22 Ma)
*** "Phlaocyon mariae"
*** "Phlaocyon marslandensis" (19 Ma)
*** "Phlaocyon minor" (30 Ma)
*** "Phlaocyon multicuspus"
*** "Phlaocyon taylori"cite journal| last = Hayes| first = F.G.| year = 2000 |title = The Brooksville 2 local fauna (Arikareean, latest Oligocene) Hernando County, Florida| journal = Bulletin of the Florida Museum of Natural History| volume = 43| issue = 1| pages= 1–47]
*** "Phlaocyon yakolai" (19 Ma)
** "Genus Protepicyon" (16 Ma)
*** "Protepicyon raki" (16 Ma)
** "Genus Protomarctus" (18 Ma)
*** "Protomarctus optatus" (18 Ma)
** "Genus Psalidocyon" (16 Ma)
*** "Psalidocyon marianae" (16 Ma)
** "Genus Rhizocyon" (30 Ma)
*** "Rhizocyon oregonensis" (30 Ma)
** "Genus Tephrocyon" (16 Ma)
*** "Tephrocyon rurestris" (16 Ma)
** "Genus Tomarctus" (16 Ma)
*** "Tomarctus brevirostris" (16 Ma)
*** "Tomarctus hippophaga" (16 Ma)

Hesperocyoninae : † (Ma = million years ago)
** "Genus Cynodesmus" (32-29 Ma)
*** "Cynodesmus martini" (29 Ma)
*** "Cynodesmus thooides" (32 Ma)
** "?Genus Caedocyon"
*** "Caedocyon tedfordi"
** "Genus Ectopocynus" (32-19 Ma)
*** "Ectopocynus antiquus" (32 Ma)
*** "Ectopocynus intermedius" (29 Ma)
*** "Ectopocynus siplicidens" (19 Ma)
** "Genus Enhydrocyon" (29-25 Ma)
*** "Enhydrocyon basilatus" (25 Ma)
*** "Enhydrocyon crassidens" (25 Ma)
*** "Enhydrocyon pahinsintewkpa" (29 Ma)
*** "Enhydrocyon stenocephalus" (29 Ma)
** "Genus Hesperocyon" (39.74-34 Ma)
*** "Hesperocyon coloradensis"
*** "Hesperocyon gregarius" (37 Ma)
** "Genus Mesocyon" (34-29 Ma)
*** "Mesocyon brachyops" (29 Ma)
*** "Mesocyon coryphaeus" (29 Ma)
*** "Mesocyn temnodon"
** "Genus Osbornodon" (32-18 Ma)
*** "Osbornodon brachypus"
*** "Osbornodon fricki" (18 Ma)
*** "Osbornodon iamonensis" (21 Ma)
*** "Osbornodon renjiei" (33 Ma)
*** "Osbornodon scitulus"cite journal| last = Wang| first = Xiaoming| year = 2003 |title = New Material of "Osbornodon" from the Early Hemingfordian of Nebraska and Florida| journal = Bulletin of the American Museum of Natural History| volume = 279| issue = | pages= 163–176| url = http://digitallibrary.amnh.org/dspace/bitstream/2246/447/19/B279a08.pdf]
*** "Osbornodon sesnoni" (32 Ma)
*** "Osbornodon wangi"
** "Genus Paraenhydrocyon" (30-25 Ma)
*** "Paraenhydrocyon josephi" (30 Ma)
*** "Paraenhydrocyon robustus" (25 Ma)
*** "Paraenhydrocyon wallovianus" (26 Ma)
** "Genus Philotrox" (29 Ma)
*** "Philotrox condoni" (29 Ma)
** "Genus Prohesperocyon" (36 Ma)
*** "Prohesperocyon wilsoni" (36 Ma)
** "Genus Sunkahetanka" (29 Ma)
*** "Sunkahetanka geringensis" (29 Ma)

ee also

* Canine reproduction

References

General references

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

* [http://www.awdconservancy.org/ African Wild Dog Conservancy]