View source for Canidae

{{Use dmy dates|date=June 2020}}
{{short description|Family of mammals}}
{{good article}}
{{Automatic taxobox
| display_parents = 3
| taxon = Canidae
| authority = [[Johann Fischer von Waldheim|Fischer de Waldheim]], 1817<ref>Fischer de Waldheim, G. 1817. Adversaria zoological. Memoir Societe Naturelle (Moscow) 5:368–428. p372</ref>
| name = Canids
| fossil_range = [[Late Eocene]]-[[Holocene]]<ref name=tedford2009/>{{rp|7}}<br />~{{fossil range|37.8|0}}
| image = <imagemap>
File:Familia Canidae.jpg|220px
rect 0 0 299 255 [[Canis]]
rect 300 0 599 255 [[Cuon]]
rect 0 256 299 488 [[Lycaon (genus)|Lycaon]]
rect 300 256 599 488 [[Cerdocyon]]
rect 0 489 299 683 [[Chrysocyon]]
rect 300 489 599 683 [[Speothos]]
rect 0 684 299 878 [[Vulpes]]
rect 300 684 599 878 [[Nyctereutes]]
rect 0 879 299 1118 [[Otocyon]]
rect 300 879 599 1118 [[Urocyon]]
</imagemap>
| image_caption = 10 of the 13 extant canid genera
| type_genus = ''[[Canis]]''
| type_genus_authority = [[Carl Linnaeus|Linnaeus]], 1758
| subdivision_ranks = Subfamilies and genera
| subdivision = *{{extinct}}''[[Prohesperocyon]]''
*{{extinct}}[[Hesperocyoninae]] 
*{{extinct}}[[Borophaginae]]
*[[Caninae]]
}}

'''Canidae''' ({{IPAc-en|ˈ|k|æ|n|ᵻ|d|iː}};<ref>Canidae. Dictionary.com. The American Heritage Stedman's Medical Dictionary. Houghton Mifflin Company. http://dictionary.reference.com/browse/Canidae (accessed: 16 February 2009).</ref> from Latin, ''[[canis]]'', "[[dog]]") is a [[family (biology)|biological family]] of caniform [[carnivora]]ns, constituting a [[clade]]. A member of this family is also called a '''canid''' ({{IPAc-en|ˈ|k|ei|n|I|d}}).{{sfn|Wang|Tedford|2008|pp=181}} The family includes three [[subfamily|subfamilies]]: the [[Caninae]], and the extinct [[Borophaginae]] and [[Hesperocyoninae]].<ref name=miklosi2015/> The Caninae are known as canines,{{sfn|Wang|Tedford|2008|pp=182}} and include [[Dog|domestic dog]]s, [[Wolf|wolves]], [[coyote]]s, [[raccoon dog]]s, [[Fox|foxes]], [[jackal]]s and other species.

Canids are found on all continents except [[Antarctica]], having arrived independently or accompanied by [[Human|human beings]] over extended periods of time. Canids vary in size from the {{Convert|2|m|ft|-long|adj=mid}}{{Citation needed|date=January 2024|reason=Article/Lemma Wolf says 160cm}} gray wolf to the {{Convert|24|cm|in|-long|adj=mid}}{{Citation needed|date=January 2024|reason=Lemma Fennec fox says 34.5cm}} [[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 [[olfactic communication|scent signals]] and vocalizations. One canid, the domestic dog, [[Origin of the domestic dog|originated]] from a [[symbiotic]] relationship with [[Upper Paleolithic]] humans and is one of the most widely kept [[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.<ref name=miklosi2015/> This subfamily includes all living canids and their most recent fossil relatives.<ref name=tedford2009/> All living canids as a group form a dental [[monophyletic]] relationship with the extinct borophagines, with both groups having a [[bicuspid]] (two points) on the lower [[carnassial]] [[talonid]], which gives this tooth an additional ability in [[mastication]]. This, together with the development of a distinct [[Glossary of mammalian dental topography#Lower teeth|entoconid]] cusp and the broadening of the talonid of the first lower [[molar (tooth)|molar]], and the corresponding enlargement of the talon of the upper first molar and reduction of its [[Glossary of mammalian dental topography#Upper teeth|parastyle]] distinguish these late [[Cenozoic]] canids and are the essential differences that identify their clade.<ref name=tedford2009/>{{rp|p6}}

The cat-like [[Feliformia]] and dog-like [[Caniformia]] emerged within the [[Carnivoramorpha]] around 45–42 Mya (million years ago).<ref>{{cite journal | last1 = Flynn | first1 = John J. | last2 = Wesley-Hunt | first2 = Gina D. | year = 2005 | title = Phylogeny of the Carnivora: Basal Relationships Among the Carnivoramorphans, and Assessment of the Position of 'Miacoidea' Relative to Carnivora | journal = Journal of Systematic Palaeontology | volume = 3 | issue = 1 | pages = 1–28 | doi=10.1017/s1477201904001518| bibcode = 2005JSPal...3....1W | s2cid = 86755875 }}</ref> 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]].<ref name=tedford2009/>{{rp|7}}

===Phylogenetic relationships===
This [[cladogram]] shows the phylogenetic position of canids within [[Caniformia]], based on fossil finds:<ref name=tedford2009/>
{{clade
 |style=font-size:90%;line-height:85%
 |label1=[[Caniformia]]
 |1={{clade
    |label1=[[Arctoidea]]
    |1= (bears, procyonids, pinnipeds, etc.) [[File:Ailurus fulgens - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|80 px]]
    |label2=[[Cynoidea]]
    |2={{clade
       |label1={{extinct}}''[[Miacis]]''
       |1=(various {{extinct}}''[[Miacis]]'' [[spp.]])[[File:Miacis restoration.jpg|60 px]]
       |state1=double
       |label2='''Canidae'''
       |2={{clade
          |label1={{extinct}}[[Hesperocyoninae]]
          |1=(''[[Mesocyon]]'', ''[[Enhydrocyon]]'', ''[[Hesperocyon]]'', and other basal canines, e.g. ''[[Cynodictis]]'')[[File:Cynodictis (white background).jpg|80 px]]
          |label2=
          |3={{clade
             |label1={{extinct}}[[Borophaginae]]
             |1=(''[[Aelurodon]]'', [[Cynarctina|cynarctin]]s, and others)<span style="{{MirrorH}}">[[File:Epicyon haydeni 3.jpg|90 px]]</span>
             |label2=[[Caninae]]
             |2= (all modern canids and extinct relatives) [[File:Dogs, jackals, wolves, and foxes (Plate X).jpg|70 px]]
             }}
            }}
        }}
     }}
 }}

==Evolution==
{{see also|Caninae#Phylogenetic relationships}}
[[File:Hesperocyoninae, Borophaginae & Caninae.jpg|thumb|right|Representatives of three canid subfamilies: ''[[Hesperocyon]]'' (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 [[Morphology (biology)|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 zone]]s exist.<ref name=Wayne>{{cite web |url=http://wooferhouse.net/Links/MolecularEvolutionOfTheDogFamily/MolecularEvolutionOfTheDogFamily.htm |title=Molecular evolution of the dog family |author=Wayne, Robert K. |access-date=27 May 2014}}</ref>

===Eocene epoch===
[[Carnivora]]ns evolved after the [[Cretaceous-Paleogene extinction event|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 [[Tympanic part of the temporal bone|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]]s.<ref name=Wang-Tedford>{{cite magazine| last = Wang| first = Xiaoming| year = 2008| title = How Dogs Came to Run the World|magazine=Natural History Magazine| volume = July/August| url = http://www.naturalhistorymag.com/features/15771/how-dogs-came-to-run-the-world| access-date = 24 May 2014 }}</ref>

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 [[Cope's rule|increase in body mass with time]] and some exhibited specialized [[hypercarnivorous]] diets that made them prone to extinction.<ref>{{cite journal | first1 = B.| last2 = Wang| last1 = Van Valkenburgh | first2 = X. | first3 = J. | title=Cope's Rule, Hypercarnivory, and Extinction in North American Canids| last3 = Damuth | s2cid = 12017658| journal=Science | volume=306| issue = #5693 | pages=101–104 | date=October 2004 | issn = 0036-8075| pmid = 15459388 | doi=10.1126/science.1102417|bibcode = 2004Sci...306..101V }}</ref>{{rp|Fig. 1}}

===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 [[viverrid]]s 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 [[evolutionary radiation|radiations]].<ref name=Gittleman>Martin, L.D. 1989. Fossil history of the terrestrial carnivora. Pages&nbsp;536–568 in J.L. Gittleman, editor. Carnivore Behavior, Ecology, and Evolution, Vol. 1. Comstock Publishing Associates: Ithaca.</ref>

===Miocene epoch===
Around 8 Mya, the [[Beringia]]n land bridge allowed members of the genus ''[[Eucyon]]'' a means to enter Asia from North America and they continued on to colonize Europe.<ref name=Perini>{{cite journal |author1=Perini, F. A. |author2=Russo, C. A. M. |author3=Schrago, C. G. |year=2010 |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1420-9101.2009.01901.x|title=The evolution of South American endemic canids: a history of rapid diversification and morphological parallelism |journal=Journal of Evolutionary Biology |volume=23 |issue=#2 |pages=311–322 |doi=10.1111/j.1420-9101.2009.01901.x |pmid=20002250|s2cid=20763999 |url-access=subscription }}</ref>

===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 [[carnassial]]s that were capable of both [[mastication]] and shearing.<ref name=Gittleman/> Around 5 million years ago, some of the Old World ''[[Eucyon]]'' evolved into the first members of ''Canis'',<ref>{{Cite journal|last1=Perri|first1=Angela R.|last2=Mitchell|first2=Kieren J.|last3=Mouton|first3=Alice|last4=Álvarez-Carretero|first4=Sandra|last5=Hulme-Beaman|first5=Ardern|last6=Haile|first6=James|last7=Jamieson|first7=Alexandra|last8=Meachen|first8=Julie|last9=Lin|first9=Audrey T.|last10=Schubert|first10=Blaine W.|last11=Ameen|first11=Carly|date=2021-01-13|title=Dire wolves were the last of an ancient New World canid lineage|url=https://www.nature.com/articles/s41586-020-03082-x|journal=Nature|volume=591|issue=7848|language=en|pages=87–91|doi=10.1038/s41586-020-03082-x|pmid=33442059|bibcode=2021Natur.591...87P|s2cid=231604957 |issn=1476-4687}}</ref> 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''.<ref>Nowak, R.M. 1979. North American Quaternary Canis. Monograph of the Museum of Natural History, University of Kansas 6:1 – 154.</ref>

The formation of the [[Isthmus of Panama]], about 3 Mya, joined [[South America]] to North America, allowing canids to [[Great American Interchange|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 fox]]es (''Lycalopex'' [[spp.]]). The monophyly of this group has been established by molecular means.<ref name=Perini/>

===Pleistocene epoch===
During the [[Pleistocene]], the North American wolf line appeared, with ''[[Canis edwardii]]'', clearly identifiable as a wolf, and ''[[Canis rufus]]'' appeared, possibly a direct descendant of ''C.&nbsp;edwardii''. Around 0.8&nbsp;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.<ref name=Larson>{{cite web |author=Larson, Robert |title=Wolves, coyotes and dogs (Genus ''Canis'') |series=The Midwestern United States 16,000&nbsp;years ago |publisher=Illinois State Museum |url=http://exhibits.museum.state.il.us/exhibits/larson/canis.html |access-date=7 June 2014}}</ref>

By 0.3 Mya, a number of subspecies of the gray wolf (''C.&nbsp;lupus'') had developed and had spread throughout Europe and northern Asia.<ref>{{cite journal |author=Nowak, R. |year=1992 |title=Wolves: The great travelers of evolution |journal=International Wolf |volume=2 |issue=4 |pages=3–7}}</ref> 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.<ref name=Chambers>{{cite journal |author1=Chambers, S.M. |author2=Fain, S.R. |author3=Fazio, B. |author4=Amaral, M. |year=2012 |title=An account of the taxonomy of North American wolves from morphological and genetic analyses |journal=North American Fauna |volume=77 |pages=1–67 |doi=10.3996/nafa.77.0001 |doi-access=free }}</ref> MtDNA studies have shown that there are at least four extant ''C. lupus'' lineages.<ref name="gaubert">{{cite journal|author1=Gaubert, P. |author2=Bloch, C. |author3=Benyacoub, S. |author4=Abdelhamid, A. |author5=Pagani, P. |display-authors=etal |year=2012 |title= Reviving the African Wolf ''Canis lupus lupaster'' in north and west Africa: A mitochondrial lineage ranging more than 6,000&nbsp;km wide |journal=PLOS ONE |volume=7 |issue=8 |page=e42740 |doi=10.1371/journal.pone.0042740 |pmid=22900047 |pmc=3416759 |bibcode=2012PLoSO...742740G|doi-access=free }}</ref> The dire wolf shared its habitat with the gray wolf, but became extinct in a large-scale extinction event that occurred around 11,500&nbsp;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.<ref name=Larson/>

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&nbsp;[[MYA (unit)|Mya]], but the African golden wolf separated 1.3&nbsp;Mya. Mitochondrial genome sequences indicated the Ethiopian wolf diverged from the wolf/coyote lineage slightly prior to that.<ref name=Koepfli-2015>{{cite journal  |last1=Koepfli |first1=Klaus-Peter |last2=Pollinger |first2=John |last3=Godinho |first3=Raquel |last4=Robinson |first4=Jacqueline |last5=Lea |first5=Amanda |last6=Hendricks |first6=Sarah |last7=Schweizer |first7=Rena M. |last8=Thalmann |first8=Olaf |last9=Silva |first9=Pedro |last10=Fan |first10=Zhenxin |last11=Yurchenko |first11=Andrey A. |last12=Dobrynin |first12=Pavel |last13=Makunin |first13=Alexey |last14=Cahill |first14=James A. |last15=Shapiro |first15=Beth |last16=Álvares |first16=Francisco |last17=Brito |first17=José C. |last18=Geffen |first18=Eli |last19=Leonard |first19=Jennifer A. |last20=Helgen |first20=Kristofer M. |last21=Johnson |first21=Warren E. |last22=o'Brien |first22=Stephen J. |last23=van Valkenburgh |first23=Blaire |last24=Wayne |first24=Robert K. |display-authors=6  |year=2015 |title=Genome-wide evidence reveals that African and Eurasian golden jackals are distinct species |journal=Current Biology |volume=25 |issue=16 |pages=2158–2165  |pmid=26234211 |doi=10.1016/j.cub.2015.06.060 |doi-access=free|bibcode=2015CBio...25.2158K }}</ref>{{rp|S1}}

=={{visible anchor|Characteristics|Anatomy}}==
{{multiple image|direction=vertical|image1=|caption1=Comparative illustration of the paws of the [[wolf]], [[golden jackal]], and [[dhole]] by A. N. Komarov|
|image2=Black-backed Jackal skeleton.jpg|caption2=Skeleton of a [[black-backed jackal]] (''Lupulella mesomelas'') on display at the [[Museum of Osteology]]
|image3=Wolf cranium labelled.jpg|caption3=Diagram of a wolf skull with key features labelled
|image4=Lupocranio.jpg|caption4=[[Eurasian wolf]] skull}}

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 [[grassland]]s. They vary in size from the fennec fox, which may be as little as {{cvt|24|cm}} in length and weigh {{cvt|0.6|kg}},<ref>{{cite book|author=Marc Tyler Nobleman|title=Foxes|url=https://books.google.com/books?id=bwZx2t3D4ukC|year=2007|publisher=Marshall Cavendish|isbn=978-0-7614-2237-2 |page=35}}</ref> to the gray [[wolf]], which may be up to {{cvt|160|cm}} long, and can weigh up to {{cvt|79|kg}}.<ref name="heptner1998">Heptner, V. G.; Naumov, N. P. (1998), [https://archive.org/details/mammalsofsov211998gept Mammals of the Soviet Union Vol. II Part 1a, Sirenia and Carnivora (Sea cows; Wolves and Bears)], Science Publishers, Inc. US., pp.&nbsp;166–176, {{ISBN|1-886106-81-9}}</ref> Only a few species are [[arboreal locomotion|arboreal]]—the [[gray fox]], the closely related [[island fox]]<ref>{{cite web|url=http://animaldiversity.org/site/accounts/information/Urocyon_littoralis.html |title=ADW: ''Urocyon littoralis'': Information |publisher=Animaldiversity.org |date=28 November 1999 |access-date=12 June 2012}}</ref> and the [[raccoon dog]] habitually climb trees.<ref>Kauhala, K.; Saeki, M. (2004). Raccoon Dog«. Canid Species Accounts. IUCN/SSC Canid Specialist Group. Pridobljeno 15 April 2009.</ref><ref>{{cite journal | title = Old dogs, new tricks: Asia's raccoon dog, a venerable member of the canid family is pushing into new frontiers | author = Ikeda, Hiroshi | journal = Natural History | volume = 95 | issue = #8 |pages=40, 44 | date = August 1986}}</ref><ref>{{Cite web |url=http://www.waza.org/en/zoo/visit-the-zoo/dogs-and-hyenas/nyctereutes-procyonoides. |title=Raccoon dog – ''Nyctereutes procyonoides''. WAZA – World Association of Zoos and Aquariums. |access-date=30 August 2011 |archive-url=https://web.archive.org/web/20150410065554/http://www.waza.org/en/zoo/visit-the-zoo/dogs-and-hyenas/nyctereutes-procyonoides |archive-date=10 April 2015 |url-status=dead }}</ref>

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 arch]]es 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.<ref name=Mivart>{{cite book |title=Dogs, Jackals, Wolves, and Foxes: A Monograph of the Canidae |last=Mivart |first=St. George Jackson |year=1890 |pages=xiv–xxxvi |url=https://archive.org/details/dogsjackalswolv00keulgoog |publisher=London : R.H. Porter : Dulau }}</ref> Females have three to seven pairs of [[mammae]].<ref name="Nowak2005">{{cite book|author=Ronald M. Nowak|title=Walker's Carnivores of the World|url=https://books.google.com/books?id=0q5aHw2mFi8C|year=2005|publisher=JHU Press|isbn=978-0-8018-8032-2}}</ref>

All canids are [[digitigrade]], meaning they walk on their toes. The [[Rhinarium|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<ref name=Mivart/> while other species have sharper, partially-retractile claws.{{cn|date=February 2023}}

The [[canine penis]] contains a [[baculum]] and a structure called the [[bulbus glandis]] that expands during [[copulation (zoology)|copulation]], forming a [[copulatory tie]] that lasts for up to an hour.<ref name="Ewer1973">{{cite book|author=Ewer, R. F. |title=The Carnivores|url=https://books.google.com/books?id=IETMd3-lSlkC|year=1973|publisher=Cornell University Press|isbn=978-0-8014-8493-3}}</ref> Young canids are born blind, with their eyes opening a few weeks after birth.<ref>{{cite book|author= Macdonald, D.|year= 1984|title= The Encyclopedia of Mammals|publisher= Facts on File|location= New York|page= [https://archive.org/details/encyclopediaofma00mals_0/page/57 57]|isbn= 978-0-87196-871-5|url= https://archive.org/details/encyclopediaofma00mals_0/page/57}}</ref> 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.{{sfn|Wang|Tedford|2008|pp=97–98}}

===Dentition===

[[Dentition]] relates to the arrangement of teeth in the mouth, with the [[Dentition#Dental formula|dental notation]] for the upper-jaw teeth using the upper-case letters I to denote [[incisors]], C for [[Canine tooth|canines]], P for [[premolars]], and M for [[molars]], and the lower-case letters i, c, p and m to denote the [[Mandible|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.{{sfn|Wang|Tedford|2008|pp=74}}

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 [[Omnivore|omnivorous]] diet.<ref name=sansalone2015/><ref name=cherin2013/> Because of its low variability, the length of the lower carnassial is used to provide an estimate of a carnivore's body size.<ref name=sansalone2015/>

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 <!-- (in [[Newton (unit)|Newtons]]/kilogram of body weight) --> 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.<ref name=wroe2005/>

Most canids have 42 [[teeth]], with a [[dentition|dental formula]] of: {{DentalFormula|upper=3.1.4.2|lower=3.1.4.3}}. 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.<ref name=Mivart/> The molar teeth are strong in most species, allowing the animals to crack open bone to reach the [[bone marrow|marrow]]. The deciduous, or baby teeth, formula in canids is {{DentalFormula|upper=3.1.3|lower=3.1.3}}, molars being completely absent.<ref name=Mivart/>

==Life history==
===Social behavior===
{{multiple image|perrow=1|image1=2012-bandipur-dhole-sambar.jpg|caption1=[[Dhole]]s attacking a [[Sambar deer|sambar]], [[Bandipur National Park]]|
|image2=Sri Lankan jackals (Canis aureus naria) male and pregnant female 5.jpg|caption2=Sri Lankan jackals [[social grooming|grooming each other]]}}
{{See also|Wolf#Behaviour|Dog behavior|Red fox#Behaviour|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.<ref>{{cite book |last1=Harris |first1=Stephen |last2=Yalden |first2=Derek |title=Mammals of the British Isles|publisher=Mammal Society|edition=4th revised |year=2008 |isbn=978-0-906282-65-6 |page=413 }}</ref> In most foxes, and in many of the true dogs, a [[breeding pair|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 [[pack (canine)|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.<ref>{{cite web |url=https://www.patriciamcconnell.com/theotherendoftheleash/comparative-canid-behavior |title=Comparative canid behaviour |author=McConnell, Patricia B. |date=31 August 2009 |work=The other end of the leash |access-date=12 June 2014}}</ref> 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.<ref name=ADW>{{cite web | title = Canidae: Coyotes, dogs, foxes, jackals, and wolves | work = Animal Diversity Web | publisher =University of Michigan| url = http://animaldiversity.org/accounts/Canidae/ | access-date = 13 June 2014}}</ref>

=== Communication ===
{{see also|Dog communication|Wolf communication}}
[[File:Red_Fox_(Vulpes_vulpes)_(W1CDR0001529_BD12).ogg|left|thumb|Red foxes barking in Pinbury Park, Gloucestershire, England.]]
Canids communicate with each other by [[Territory (animal)|scent signals]], by visual clues and gestures, and by vocalizations such as growls, barks, and [[Howl (sound)|howls]]. In most cases, groups have a home territory from which they drive out other [[conspecific]]s. Canids use [[Urine marking|urine scent marks]]<ref>{{Cite book |last=Doty |first=Richard |url=https://books.google.com/books?id=PWwCnplyxYYC&pg=PA284 |title=Mammalian Olfaction, Reproductive Processes, and Behavior |date=2012-12-02 |publisher=Elsevier |isbn=978-0-323-15450-5 |language=en}}</ref><ref>{{Cite book |last=Estes |first=Richard |url=https://books.google.com/books?id=g977LsZHpcsC&dq=canid&pg=PA387 |title=The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates |date=1991 |publisher=University of California Press |isbn=978-0-520-08085-0 |language=en}}</ref> to mark their [[Cache (biology)|food cache]]s<ref>{{Cite book |last=Muller-Schwarze |first=Dietland |url=https://books.google.com/books?id=HaaFlUw4goIC&pg=PA167 |title=Chemical Ecology of Vertebrates |date=2006-09-07 |publisher=Cambridge University Press |isbn=978-1-139-45730-9 |language=en}}</ref> or warn trespassing individuals.<ref>Nowak, R. M.; Paradiso, J. L. 1983. ''Walker's Mammals of the World''. Baltimore, Maryland: The Johns Hopkins University Press. {{ISBN|0-8018-2525-3}}.</ref> Social behavior is also mediated by secretions from glands on the upper surface of the tail near its root and from the [[anal gland]]s,<ref name=ADW/> [[preputial gland]]s,<ref>{{Cite book |last1=Mech |first1=L. David |url=https://books.google.com/books?id=_mXHuSSbiGgC&pg=PA82 |title=Wolves: Behavior, Ecology, and Conservation |last2=Boitani |first2=Luigi |date=2010-10-01 |publisher=University of Chicago Press |isbn=978-0-226-51698-1 |language=en}}</ref><ref>Van Heerden, Joseph. "[https://repository.up.ac.za/bitstream/handle/2263/51834/5vanheerden1981.pdf?sequence=1 The role of integumental glands in the social and mating behaviour of the hunting dog Lycaon pictus (Temminck, 1820)]." (1981).</ref> and [[supracaudal gland]]s.<ref>Fox, Michael W., and James A. Cohen. "[https://publish.iupress.indiana.edu/read/how-animals-communicate/section/d60e31a7-f9c0-4f0d-b569-7a922f64e0c1 Canid communication]." How animals communicate (1977): 728-748.</ref>


===Reproduction===
{{See also|Canine reproduction}}
{{image frame|border=no|content={{photo montage|size=220
|photo1a=Korean wolves mating (cropped).jpg
|photo1b=Red foxes mating (2).jpg
|border=0
|color=transparent
|text=Mating [[Korean wolf|Korean wolves]] and [[red fox]]es
}}
{{photo montage|size=220
|photo1a=Wilde huendin am stillen.jpg
|border=0
|color=transparent
|text=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 [[Monogamy in animals|monogamous]], provide paternal care to their offspring, have reproductive cycles with lengthy [[Estrous cycle#Proestrus|proestral]] and [[Estrous cycle#Metestrus or diestrus|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 [[alloparent]]al care they can provide to help raise the next generation.<ref name=Asa>{{cite book |title=A review of Small Canid Reproduction: in The Swift Fox: Ecology and Conservation of Swift Foxes in a Changing World |editor-last=Asa |editor-first=Cheryl S. |editor2-last=Valdespino|editor2-first=Carolina|editor3-last=Carbyn|editor3-first=Ludwig N.|editor4-last= Sovada|editor4-first=Marsha Ann |year=2003 |publisher=University of Regina Press |isbn=978-0-88977-154-3 |pages=117–123 |url=https://books.google.com/books?id=Er1di1OqnlAC&pg=PA117}}</ref> Most canid species are spontaneous ovulators,<ref>{{Cite book |last=Dixson |first=Alan F. |url=https://books.google.com/books?id=whcvEAAAQBAJ&q=canid |title=Mammalian Sexuality: The Act of Mating and the Evolution of Reproduction |date=2021-06-03 |publisher=Cambridge University Press |isbn=978-1-108-69949-5 |language=en}}</ref> though [[maned wolves]] are [[induced ovulation (animals)|induced ovulators]].<ref>{{Cite book |last1=Consorte-McCrea |first1=Adriana G. |url=https://books.google.com/books?id=zT3SBQAAQBAJ&dq=canid+%22induced%22+ovulation&pg=PA106 |title=Ecology and Conservation of the Maned Wolf: Multidisciplinary Perspectives |last2=Santos |first2=Eliana Ferraz |date=2013-10-24 |publisher=CRC Press |isbn=978-1-4665-1260-3 |language=en}}</ref> 

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 [[Estrous cycle#Anestrus|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.<ref name=Asa/> Canids have an oestrus period of 1 to 20 days, lasting one week in most species.<ref>{{Cite book |last1=Solomon |first1=Nancy G. |url=https://books.google.com/books?id=6_brrHFGmEcC&pg=PA139 |title=Cooperative Breeding in Mammals |last2=French |first2=Jeffrey A. |date=1997-03-13 |publisher=Cambridge University Press |isbn=978-0-521-45491-9 |language=en}}</ref>

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.<ref name=Mivart/> 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.<ref>{{cite book |title=The Behavioural Biology of Dogs |editor-last=Jensen |editor-first=Per |year=2007 |publisher=CABI |isbn=978-1-84593-188-9 |pages=158–159 |url=https://books.google.com/books?id=SpkSd__EdKYC&pg=PA3 }}</ref> Young canids may take a year to mature and learn the skills they need to survive.<ref>Voelker, W. 1986. ''The Natural History of Living Mammals''. Medford, New Jersey: Plexus Publishing. {{ISBN|0-937548-08-1}}</ref> 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.<ref>{{cite web |url=http://www.animalinfo.org/species/carnivor/lycapict.htm |title=''Lycaon pictus'' |date=26 November 2005 |work=Animal Info: Endangered animals of the world |access-date=11 June 2014}}</ref>

==Canids and humans==
[[File:BedaleHunt2005.jpg|thumb|Traditional English [[fox hunt]]]]
{{further|Dog}}

One canid, the [[Origin of the domestic dog|domestic dog]], entered into a partnership with humans a long time ago. The dog was the first domesticated species.<ref name=larson2014/><ref name=larson2012/><ref name=britannica/><ref name=perri2016/> The archaeological record shows the first undisputed dog remains buried beside humans 14,700&nbsp;years ago,<ref name=giemsch2015/> with disputed remains occurring 36,000&nbsp;years ago.<ref name=germonpre2009/> These dates imply that the earliest dogs arose in the time of human [[hunter-gatherer]]s and not [[agriculturists]].<ref name=thalmann2013>{{Cite journal|author=Thalmann, O.|year=2013|title=Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs|journal=Science|doi=10.1126/science.1243650|pmid=24233726|volume=342|issue=#6160|pages=871–4|bibcode=2013Sci...342..871T|s2cid=1526260|url=https://digital.csic.es/bitstream/10261/88173/1/accesoRestringido.pdf |archive-url=https://web.archive.org/web/20140104071156/http://digital.csic.es/bitstream/10261/88173/1/accesoRestringido.pdf |archive-date=2014-01-04 |url-status=live}}</ref><ref name=freedman2014/>

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.<ref>{{cite journal|author1=Schleidt, Wolfgang M. |author2=Shalter, Michael D. |year=2003 |title=Co-evolution of humans and canids: An alternative view of dog domestication: Homo homini lupus? |journal=Evolution and Cognition |volume=9 |issue=1 |pages=57–72 |url=https://www.uwsp.edu/psych/s/275/Science/Coevolution03.pdf |url-status=dead |archive-url=https://web.archive.org/web/20140411142919/http://www4.uwsp.edu/psych/s/275/science/coevolution03.pdf |archive-date=11 April 2014 }}</ref>

Among canids, only the gray wolf has widely been known to prey on humans.<ref>{{cite book |author=Kruuk, H. |year=2002 |title=Hunter and Hunted: Relationships between carnivores and people |location=Cambridge, UK |publisher=Cambridge University Press |isbn=0-521-81410-3}}</ref>{{pn|date=November 2021}} Nonetheless, at least [[Coyote attack|two records]] of coyotes killing humans have been published,<ref name="AOH">{{cite web |title=Coyote attacks: An increasing suburban problem |publisher=San Diego County, California |url=http://www.co.san-diego.ca.us/awm/docs/coyoteattacks.pdf |access-date=19 August 2007 |archive-url=https://web.archive.org/web/20060224153601/http://www.co.san-diego.ca.us/awm/docs/coyoteattacks.pdf |archive-date=24 February 2006 }}</ref> and at least two other reports of golden jackals killing children.<ref name="CWC">{{cite web |title=''Canis aureus'' |website=Animal Diversity Web |publisher=University of Michigan |url=http://animaldiversity.org/site/accounts/information/Canis_aureus.html |access-date=31 July 2007 }}</ref> 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.<ref>{{cite news |title=Fox hunting worldwide |url=http://news.bbc.co.uk/1/hi/uk/428122.stm |newspaper=BBC News |date=16 September 1999 |access-date=16 June 2014 }}</ref> Canids such as the [[dhole]] are now [[Endangered species|endangered]] in the wild because of persecution, habitat loss, a depletion of ungulate prey species and transmission of diseases from domestic dogs.<ref>{{cite iucn |author=Kamler, J.F. |author2=Songsasen, N. |author3=Jenks, K. |author4=Srivathsa, A. |author5=Sheng, L. |author6=Kunkel, K. |date=2015 |title=''Cuon alpinus'' |volume=2015 |page=e.T5953A72477893 |doi=10.2305/IUCN.UK.2015-4.RLTS.T5953A72477893.en |access-date=11 November 2021}}</ref>

==See also==

*[[List of canids]]
*[[Largest wild canids]]
*[[Canid hybrid]]
*[[Free-ranging dog]]

==References==<!-- SystBioi46:622. ZoolScripta33:311. -->
{{Reflist|30em|refs=

<ref name=britannica>{{cite encyclopedia|url=https://www.britannica.com/science/domestication|access-date=26 May 2016|encyclopedia=Encyclopædia Britannica|title=Domestication|year=2016}}</ref>

<ref name=cherin2013>{{cite journal|last1=Cherin|first1=Marco|last2=Bertè|first2=Davide Federico |last3=Sardella|first3=Raffaele|last4=Rook|first4=Lorenzo|title=Canis etruscus (Canidae, Mammalia) and its role in the faunal assemblage from Pantalla (Perugia, central Italy): comparison with the Late Villafranchian large carnivore guild of Italy|journal=Bollettino della Società Paleontologica Italiana|volume=52|issue=#1|year=2013|pages=11–18|url=https://www.researchgate.net/publication/256431568}}</ref>

<ref name=freedman2014>{{Cite journal|author=Freedman, A.|year=2014|title=Genome sequencing highlights the dynamic early history of dogs|journal=[[PLOS Genetics]]|volume=10|issue=#1|pages=e1004016|doi=10.1371/journal.pgen.1004016|pmid=24453982|pmc=3894170 |doi-access=free }}</ref>

<ref name=germonpre2009>{{cite journal|author=Germonpre, M.|year=2009 |title=Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: Osteometry, ancient DNA and stable isotopes|journal=Journal of Archaeological Science |volume=36 |issue=#2 |pages=473–490 | doi = 10.1016/j.jas.2008.09.033|bibcode=2009JArSc..36..473G }}</ref>
<ref name=giemsch2015>Liane Giemsch, Susanne C. Feine, Kurt W. Alt, Qiaomei Fu, Corina Knipper, Johannes Krause, Sarah Lacy, Olaf Nehlich, Constanze Niess, Svante Pääbo, Alfred Pawlik, Michael P. Richards, Verena Schünemann, Martin Street, Olaf Thalmann, Johann Tinnes, Erik Trinkaus & Ralf W. Schmitz. "Interdisciplinary investigations of the late glacial double burial from Bonn-Oberkassel". Hugo Obermaier Society for Quaternary Research and Archaeology of the Stone Age: 57th Annual Meeting in Heidenheim, 7th – 11 April 2015, 36-37</ref>

<ref name=larson2012>{{cite journal|author=Larson G|year=2012|title=Rethinking dog domestication by integrating genetics, archeology, and biogeography|journal=PNAS|volume=109|issue=#23|pages=8878–8883|doi=10.1073/pnas.1203005109|pmid=22615366|pmc=3384140|bibcode=2012PNAS..109.8878L|doi-access=free}}</ref>
<ref name=larson2014>{{cite journal|vauthors=Larson G, Bradley DG |year=2014|title=How Much Is That in Dog Years? The Advent of Canine Population Genomics|journal=PLOS Genetics |doi=10.1371/journal.pgen.1004093|pmid=24453989|pmc=3894154|volume=10|issue=#1|pages=e1004093 |doi-access=free }}</ref>

<ref name=miklosi2015>{{cite book|last1=Miklosi|first1=Adam|title=Dog Behaviour, Evolution, and Cognition|publisher=Oxford University Press|edition=2nd|series=Oxford Biology|year=2015|pages=103–107|url=https://books.google.com/books?id=VT-WBQAAQBAJ&pg=PA103|isbn=978-0199545667}}</ref>

<ref name=perri2016>{{cite journal|doi=10.1016/j.jas.2016.02.003|title=A wolf in dog's clothing: Initial dog domestication and Pleistocene wolf variation|journal=Journal of Archaeological Science|volume=68|pages=1–4|year=2016|last1=Perri|first1=Angela|bibcode=2016JArSc..68....1P }}</ref>

<ref name=sansalone2015>{{cite journal|doi=10.1016/j.quascirev.2014.12.009|title=Evolutionary trends and stasis in carnassial teeth of European Pleistocene wolf Canis lupus (Mammalia, Canidae)|journal=Quaternary Science Reviews|volume=110|pages=36–48|year=2015|last1=Sansalone|first1=Gabriele|last2=Bertè|first2=Davide Federico|last3=Maiorino|first3=Leonardo|last4=Pandolfi|first4=Luca|hdl=11380/1318343|hdl-access=free}}</ref>

<ref name=tedford2009>{{cite journal |title=Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae) |year=2009 |first1=Richard |last1=Tedford |author-link1=Richard H. Tedford |first2=Xiaoming |last2=Wang |author-link2=Xiaoming Wang (paleontologist) |first3=Beryl E. |last3=Taylor |journal=[[Bulletin of the American Museum of Natural History]] |volume=325 |pages=1–218 |doi=10.1206/574.1 |hdl=2246/5999 |s2cid=83594819 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/5999/1/B325.pdf |archive-url=https://web.archive.org/web/20120406021726/http://digitallibrary.amnh.org/dspace/bitstream/2246/5999/1/B325.pdf |archive-date=2012-04-06 |url-status=live}}</ref>

<ref name=wroe2005>{{cite journal|doi=10.1098/rspb.2004.2986|title=Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa|journal=Proceedings of the Royal Society B: Biological Sciences|volume=272|issue=#1563|pages=619–25|year=2005|last1=Wroe|first1=S.|last2=McHenry|first2=C.|last3=Thomason|first3=J.|pmid=15817436|pmc=1564077}}</ref>
}}

==Bibliography==
* {{cite book|last1=Wang|first1=Xiaoming|author-link1=Xiaoming Wang (paleontologist)|last2=Tedford|first2=Richard H.|author-link2=Richard H. Tedford|title=Dogs:Their Fossil Relatives and Evolutionary History|publisher=Columbia University Press, New York|year=2008|isbn=978-0-231-13529-0|url={{Google books|plainurl=yes|id=ctaMZMJIf-oC}}}}

==External links==
{{Wikispecies}}
{{Commons category|Canidae}}
* {{NCBI taxid|9608}}

{{Carnivora|Ca.}}
{{Canidae extinct nav}}

{{Taxonbar|from=Q25324}}
{{Authority control}}

[[Category:Canids| ]]
[[Category:Mammal families]]
[[Category:Bartonian first appearances]]
[[Category:Extant Eocene first appearances]]
[[Category:Taxa named by Gotthelf Fischer von Waldheim]]