View source for Canis

{{Short description|Genus of carnivores}}
{{About|the genus of canines}}
{{Automatic taxobox
| fossil_range = {{fossil range|Pliocene|Present}}Possible [[Late Miocene]] origin<ref name=wang2008/>
| image = Canis.png
| image_caption = 1st row: [[wolf]] (''C. lupus''),<br />[[dog]] (''C. familiaris'');<br />2nd row: [[red wolf]] (''C. rufus''),<br />[[eastern wolf]] (''C. lycaon'');<br />3rd row: [[coyote]] (''C. latrans''),<br />[[golden jackal]] (''C. aureus'');<br />4th row: [[Ethiopian wolf]] (''C. simensis''),<br />[[African wolf]] (''C. lupaster'').
| taxon = Canis
| display_parents = 3
| authority = [[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]<ref name=linnaeus1758/>
| type_species = ''[[Canis familiaris]]''
| type_species_authority = [[Carl Linnaeus|Linnaeus]], 1758
| subdivision_ranks = Species
| subdivision = Extant:
*''[[Canis aureus]]''
*''[[Canis familiaris]]''
*''[[Canis latrans]]''
*''[[Canis lupaster]]''
*''[[Canis lupus]]''
*''[[Canis lycaon]]''
*''[[Canis rufus]]''
*''[[Canis simensis]]''
Extinct:
*† ''[[Canis adoxus]]''
*† ''[[Evolution of the wolf#Canis borjgali|Canis borjgali]]''
*† ''[[Canis chihliensis]]''
*† ''[[Canis edwardii]]''
*† ''[[Canis etruscus]]''
*† ''[[Canis lepophagus]]''
*† ''[[Canis mosbachensis]]''
*† ''[[Canis palmidens]]''{{citation needed|date=April 2023}}
*† ''[[Canis variabilis]]''
*† '''Subgenus ''[[Xenocyon]]'''''
**† ''[[Canis africanus]]''
**† ''[[Canis antonii]]''
**† ''[[Canis falconeri]]''
**† ''[[Canis lycanoides]]''
}}
'''''Canis''''' is a [[genus]] of the [[Caninae]] which includes multiple [[extant taxon|extant]] species, such as [[Wolf|wolves]], [[dog]]s, [[coyotes]], and [[golden jackals]]. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.<ref name=heptner1998>Heptner, V. G.; Naumov, N. P. (1998). ''Mammals of the Soviet Union'' Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea Cows, Wolves and Bears). Science Publishers, Inc. USA. pp. 124–129. {{ISBN|1-886106-81-9}}.</ref>

==Taxonomy==
The [[genus]] ''Canis'' ([[Carl Linnaeus]], 1758) was published in the [[10th edition of Systema Naturae]]<ref name=linnaeus1758/> and included the dog-like carnivores: the domestic dog, wolves, coyotes and jackals. All species within ''Canis'' are [[Phylogenetics|phylogenetically]] closely related with 78 [[chromosome]]s and can potentially [[hybrid (biology)|interbreed]].<ref name=wayne1999/> In 1926, the [[International Commission on Zoological Nomenclature]] (ICZN) in Opinion 91 included Genus ''Canis'' on its ''Official Lists and Indexes of Names in Zoology''.<ref name=iczn1926/> In 1955, the ICZN's Direction 22 added ''[[Dog|Canis familiaris]]'' as the [[type species]] for genus ''Canis'' to the official list.<ref name=iczn1955/>

{{Blockquote|''Canis'' is primitive relative to ''Cuon'', ''Lycaon'', and ''Xenocyon'' in its relatively larger canines and lack of such dental adaptations for hypercarnivory as m1–m2 metaconid and entoconid small or absent; M1–M2 hypocone small; M1–M2 lingual cingulum weak; M2 and m2 small, may be single-rooted; m3 small or absent; and wide palate.|[[Richard H. Tedford]]<ref name=Tedford2009/>}}

The cladogram below is based on the [[DNA]] phylogeny of Lindblad-Toh ''et al''. (2005),<ref name=LindbladToh2005/> modified to incorporate recent findings on ''Canis'' species,<ref name=Koepfli2015/><ref>{{Cite journal |last1=Wilson |first1=Paul J. |last2=Grewal |first2=Sonya |last3=Lawford |first3=Ian D. |last4=Heal |first4=Jennifer NM |last5=Granacki |first5=Angela G. |last6=Pennock |first6=David |last7=Theberge |first7=John B. |last8=Theberge |first8=Mary T. |last9=Voigt |first9=Dennis R. |last10=Waddell |first10=Will |last11=Chambers |first11=Robert E. |date=2011-02-15 |title=DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf |url=https://cdnsciencepub.com/doi/abs/10.1139/z00-158 |journal=Canadian Journal of Zoology |volume=78 |issue=12 |pages=2156–2166 |language=en |doi=10.1139/z00-158|url-access=subscription }}</ref>

{{Clade |style=font-size:85%; line-height:85%
|label1=''Canis''
|1={{Clade
   |1={{Clade
      |1={{Clade
         |1={{Clade
            |1={{Clade
               |1={{Clade 
                  |1=''[[Canis latrans]]'' (coyote) [[File:Dogs, jackals, wolves, and foxes (Plate IX).png|50 px]]      
                  |2={{Clade
                     |1=''[[Canis rufus]]'' (red wolf) [[File:Dogs, jackals, wolves, and foxes (Plate V) C. l. rufus mod.png|50 px]]
                     |2=''[[Canis lycaon]]'' (eastern wolf) [[File:Dogs, jackals, wolves, and foxes (Plate V).png|50 px]]  
                  }} }}
               |2={{Clade
                  |1=''[[Canis lupus]]'' (gray wolf) [[File:Dogs, jackals, wolves, and foxes (Plate I).png|50 px]]
                  |2=''[[Canis familiaris]]'' (domestic dog) <span style="{{MirrorH}}">[[File:202104 Dog.svg|50 px]]</span>
               }} }}
            |2=''[[Canis lupaster]]'' ([[African golden wolf]]) [[File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XI).png|50px]]
            }}
         |2=''[[Canis simensis]]'' ([[Ethiopian wolf]]) [[File:Dogs, jackals, wolves, and foxes (Plate VI).png|50 px]]
         }}
      |2=''[[Canis aureus]]'' ([[golden jackal]]) [[File:Dogs, jackals, wolves, and foxes (Plate X).png|50 px]]
}} }} }}

In 2019, a workshop hosted by the [[IUCN]]/SSC Canid Specialist Group recommends that because DNA evidence shows the [[side-striped jackal]] (''Canis adustus'') and [[black-backed jackal]] (''Canis mesomelas'') to form a monophyletic lineage that sits outside of the ''Canis''/''Cuon''/''Lycaon'' clade, that they should be placed in a distinct genus, ''Lupulella'' Hilzheimer, 1906 with the names ''Lupulella adusta'' and ''Lupulella mesomelas''.<ref name=Alvares2019/>

===Evolution===
:''See further: [[Canidae#Evolution|Evolution of the canids]]''
The fossil record shows that [[Feliformia|feliform]]s and [[Caniformia|caniform]]s emerged within the clade [[Carnivoramorpha]] 43&nbsp;million [[Years before present|YBP]].<ref name=flynn2005/> The caniforms included the fox-like genus ''[[Leptocyon]]'', whose various species existed from 24&nbsp;million YBP before branching 11.9&nbsp;million YBP into ''[[Vulpes]]'' (foxes) and Canini (canines). The jackal-sized ''[[Eucyon]]'' existed in North America from 10&nbsp;million YBP and by the [[Early Pliocene]] about 6-5&nbsp;million YBP the coyote-like ''Eucyon davisi''<ref name=fossilworks5/> invaded Eurasia. The canids that had emigrated from North America to Eurasia – ''[[Eucyon]]'', ''[[Vulpes]]'', and ''[[Nyctereutes]]'' – were small to medium-sized predators during the Late Miocene and Early Pliocene but they were not the top predators.

[[File:C. dirus, C. lupus, C. lycaon, C. rufus, C. latrans, C. anthus C. aureus & C. mesomelas skulls.jpg|thumb|Skulls of dire wolf (''Aenocyon dirus''), gray wolf (''C. lupus''), eastern wolf (''C. lycaon''), red wolf (''C. rufus''), coyote (''C. latrans''), African golden wolf (''C. lupaster''), golden jackal (''C. aureus'') and black-backed jackal (''Lupulella mesomelas'')]]

For ''Canis'' populations in the New World, ''Eucyon'' in North America gave rise to early North American ''Canis'' which first appeared in the [[Miocene]] (6&nbsp;million YBP) in south-western United States and Mexico. By 5&nbsp;million YBP the larger ''[[Canis lepophagus]]'', ancestor of wolves and coyotes, appeared in the same region.<ref name="wang2008" />{{rp|p58}}

Around 5 million years ago, some of the Old World ''Eucyon'' evolved into the first members of ''Canis'',<ref name="Perri-2021">{{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> and the position of the canids would change to become a dominant predator across the [[Palearctic realm|Palearctic]]. The wolf-sized ''[[Evolution of the wolf#Canis chihliensis|C. chihliensis]]'' appeared in northern China in the Mid-Pliocene around 4-3&nbsp;million YBP. This was followed by an explosion of ''Canis'' evolution across Eurasia in the Early Pleistocene around 1.8&nbsp;million YBP in what is commonly referred to as the ''wolf event''. It is associated with the formation of the [[mammoth steppe]] and continental glaciation. ''Canis'' spread to Europe in the forms of ''[[Canis arnensis|C. arnensis]]'', ''[[Canis etruscus|C. etruscus]]'', and ''[[Canis falconeri|C. falconeri]]''.<ref name=wang2008/>{{rp|p148}}

However, a 2021 genetic study of the [[dire wolf]] (''Aenocyon dirus''), previously considered a member of ''Canis'', found that it represented the last member of an ancient lineage of canines originally indigenous to the New World that had diverged prior to the appearance of ''Canis'', and that its lineage had been distinct since the Miocene with no evidence of introgression with ''Canis''. The study hypothesized that the [[Neogene]] canids in the New World, ''[[Armbruster's wolf|Canis armbrusteri]]'' and ''[[Canis edwardii]]'', were possibly members of the distinct dire wolf lineage that had [[Convergent evolution|convergently evolved]] a very similar appearance to members of ''Canis''. True members of ''Canis'', namely the [[Gray Wolf|gray wolf]] and [[coyote]], likely only arrived in the New World during the [[Late Pleistocene]], where their dietary flexibility and/or ability to hybridize with other canids allowed them to survive the [[Quaternary extinction event]], unlike the dire wolf.<ref name="Perri-2021" />

''[[Xenocyon]]'' (strange wolf) is an extinct [[subgenus]] of ''Canis''.<ref name=rook1994/> The diversity of the ''Canis'' group decreased by the end of the [[Early Pleistocene]] to the [[Middle Pleistocene]] and was limited in Eurasia to the small wolves of the ''[[Canis mosbachensis]]–Canis variabilis'' group and the large hypercarnivorous [[Xenocyon#Canis (Xenocyon) lycaonoides|''Canis (Xenocyon) lycaonoides'']].<ref name=sotnikova2010/> The hypercarnivore ''Xenocyon'' gave rise to the modern [[dhole]] and the [[African wild dog]].<ref name=wang2008/>{{rp|p149}}

==Dentition and biteforce==
[[File:Wolf cranium labelled.jpg|thumb|300px|Diagram of a wolf skull with key features labelled]]
[[File:Lupocranio.jpg|thumb|right|[[Eurasian wolf]] skull]]

{| class="wikitable sortable"
|+ [[Bite force quotient|Bite force]] adjusted for body weight in [[Newton (unit)|Newtons]] per kilogram<ref name=Christiansen2007/>
! scope="col" | Canid
! scope="col" | [[Carnassial]]
! scope="col" | [[Canine tooth|Canine]]  
|-
! scope="row" | Gray wolf
| 131.6
| 127.3
|-
! scope="row" | [[Dhole]]
| 130.7
| 132.0
|-
! scope="row" | [[African wild dog]]
| 127.7
| 131.1
|-
! scope="row" | [[Greenland dog]] and [[dingo]]
| 117.4
| 114.3
|-
! scope="row" | [[Coyote]]
| 107.2
| 98.9
|-
! scope="row" | [[Side-striped jackal]]
| 93.0
| 87.5
|-
! scope="row" | [[Golden jackal]]
| 89.6
| 87.7
|-
! scope="row" | [[Black-backed jackal]]
| 80.6
| 78.3
|-
|}

<!--Note: [[Dire wolf]] and [[Beringian wolf]] both link to here.-->
[[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.<ref name=wang2008/>{{rp|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 candid'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 hunting 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/>

==Behavior==

===Description and sexual dimorphism===
{{multiple image|perrow=4|total_width=500|image2=Lactating Female Coyote - cropped.jpg|image1=Coyote 05282020000047416 (49945760021).jpg
|image4=Female_Gray_Wolf_(6045671049).jpg|image3=20140812 WOLF IMG 1043.png
|caption1=Male coyote|caption2=Female coyote|caption3=Male gray wolf|caption4=Female gray wolf}}

There is little variance among male and female canids. Canids tend to live as monogamous pairs. Wolves, [[dholes]], [[coyote]]s, and [[jackals]] live in groups that include [[breeding pair]]s and their offspring. Wolves may live in extended family groups. To take prey larger than themselves, the African wild dog, the dhole, and the gray wolf depend on their jaws as they cannot use their forelimbs to grapple with prey. They work together as a pack consisting of an alpha pair and their offspring from the current and previous years.<ref name=valkenburgh2002/> Social mammal predators prey on herbivores with a body mass similar to that of the combined mass of the predator pack.<ref name=sorkin2008/><ref name=earle1987/> The gray wolf specializes in preying on the vulnerable individuals of large prey,<ref name=paquet2003/> and a pack of timber wolves can bring down a {{convert|500|kg|lb|abbr=on}} moose.<ref name=mech1966/><ref name=anyonge2006/>

==== Mating behaviour ====
The genus ''Canis'' contains many different species and has a wide range of different mating systems that varies depending on the type of canine and the species.<ref name="Current Zoology">{{Cite journal|last1=Dale|first1=Rachel|last2=Marshall-Pescini|first2=Sarah|last3=Range|first3=Friederike|date=2017-06-01|title=Do females use their sexual status to gain resource access? Investigating food-for-sex in wolves and dogs|journal=Current Zoology|language=en|volume=63|issue=3|pages=323–330|doi=10.1093/cz/zow111|pmid=29491991|pmc=5804177|issn=1674-5507}}</ref> In a study done in 2017, it was found that in some species of canids females use their sexual status to gain food resources. The study looked at wolves and dogs. [[Gray wolf|Wolves]] are typically [[Monogamy in animals|monogamous]] and form [[pair bond|pair-bonds]]; whereas dogs are promiscuous when free-range and mate with multiple individuals. The study found that in both species females tried to gain access to food more and were more successful in monopolizing a food resource when in heat. Outside of the breeding season their efforts were not as persistent or successful. This shows that the food-for-sex hypothesis likely plays a role in the food sharing among canids and acts as a direct benefit for the females.<ref name="Current Zoology"/>

Another study on [[free-ranging dog]]s found that social factors played a significant role in the determination of mating pairs. The study, done in 2014, looked at social regulation of reproduction in the dogs.<ref name="Social Variables Affecting Mate Preferences, Copulation and Reproductive Outcome in a Pack of Free-Ranging Dogs">{{Cite journal|last1=Cafazzo|first1=Simona|last2=Bonanni|first2=Roberto|last3=Valsecchi|first3=Paola|last4=Natoli|first4=Eugenia|date=2014-06-06|title=Social Variables Affecting Mate Preferences, Copulation and Reproductive Outcome in a Pack of Free-Ranging Dogs|journal=PLOS ONE|language=en|volume=9|issue=6|pages=e98594|doi=10.1371/journal.pone.0098594|pmid=24905360|pmc=4048177|bibcode=2014PLoSO...998594C|issn=1932-6203|doi-access=free}}</ref> They found that females in [[Estrus|heat]] searched out [[dominance (ethology)|dominant]] males and were more likely to mate with a dominant male who appeared to be a quality leader. The females were more likely to reject submissive males. Furthermore, cases of [[male-male competition]] were more aggressive in the presence of high ranking females. This suggests that females prefer dominant males and males prefer high ranking females meaning social cues and status play a large role in the determination of mating pairs in dogs.<ref name="Social Variables Affecting Mate Preferences, Copulation and Reproductive Outcome in a Pack of Free-Ranging Dogs"/>

Canids also show a wide range of [[parental care]] and in 2018 a study showed that [[sexual conflict]] plays a role in the determination of [[Parental investment|intersexual parental investment]].<ref>{{Cite journal|last1=Schell|first1=Christopher J|last2=Young|first2=Julie K|last3=Lonsdorf|first3=Elizabeth V|last4=Mateo|first4=Jill M|last5=Santymire|first5=Rachel M|title=It takes two: Evidence for reduced sexual conflict over parental care in a biparental canid|journal=Journal of Mammalogy|volume=99|issue=1|pages=75–88|doi=10.1093/jmammal/gyx150|year=2018|doi-access=free}}</ref> The studied looked at [[coyote]] mating pairs and found that paternal investment was increased to match or near match the maternal investment. The amount of parental care provided by the fathers also was shown to fluctuated depending on the level of care provided by the mother.

Another study on parental investment showed that in free-ranging dogs, mothers modify their energy and time investment into their pups as they age.<ref>{{Cite journal|last1=Paul|first1=Manabi|last2=Sau|first2=Shubhra|last3=Nandi|first3=Anjan K.|last4=Bhadra|first4=Anindita|date=2017-01-01|title=Clever mothers balance time and effort in parental care: a study on free-ranging dogs|journal=Royal Society Open Science|language=en|volume=4|issue=1|pages=160583|doi=10.1098/rsos.160583|pmid=28280555|pmc=5319321|issn=2054-5703|arxiv=1607.01135|bibcode=2017RSOS....460583P}}</ref> Due to the high mortality of free-range dogs at a young age a mother's fitness can be drastically reduced. This study found that as the pups aged the mother shifted from high-energy care to lower-energy care so that they can care for their offspring for a longer duration for a reduced energy requirement. By doing this the mothers increasing the likelihood of their pups surviving infancy and reaching adulthood and thereby increase their own fitness.

A study done in 2017 found that aggression between male and female gray wolves varied and changed with age.<ref>{{Cite journal|last1=Cassidy|first1=Kira A.|last2=Mech|first2=L. David|last3=MacNulty|first3=Daniel R.|last4=Stahler|first4=Daniel R.|last5=Smith|first5=Douglas W.|title=Sexually dimorphic aggression indicates male gray wolves specialize in pack defense against conspecific groups|journal=Behavioural Processes|volume=136|pages=64–72|doi=10.1016/j.beproc.2017.01.011|pmid=28143722|year=2017|s2cid=32107025|url=https://digitalcommons.unl.edu/usgsnpwrc/311|url-access=subscription}}</ref> Males were more likely to chase away rival packs and lone individuals than females and became increasingly aggressive with age. Alternatively, females were found to be less aggressive and constant in their level of aggression throughout their life. This requires further research but suggests that intersexual aggression levels in [[Wolf|gray wolves]] relates to their [[mating system]].

====Tooth breakage====
[[File:Wolf dentition in the Ice Age.svg|thumb|200px|Dentition of a wolf showing functions of the teeth.]]

<!--Note: [[Dire wolf]] and [[Beringian wolf]] both link to here.-->
Tooth breakage is a frequent result of carnivores' feeding behaviour.<ref name=valkenburgh1993a/> Carnivores include both [[pack hunter]]s and solitary hunters. The solitary hunter depends on a powerful bite at the canine teeth to subdue their prey, and thus exhibits a strong [[mandibular symphysis]]. In contrast, a pack hunter, which delivers many shallower bites, has a comparably weaker mandibular symphysis. Thus, researchers can use the strength of the mandibular symphysis in fossil carnivore specimens to determine what kind of hunter it was{{snd}}a pack hunter or a solitary hunter{{snd}}and even how it consumed its prey. The mandibles of canids are buttressed behind the carnassial teeth to crack bones with their post-carnassial teeth (molars M2 and M3). A study found that the modern gray wolf and the red wolf (''C.{{nbsp}}rufus'') possess greater buttressing than all other extant canids and the extinct dire wolf. This indicates that these are both better adapted for cracking bone than other canids.<ref name=therrien2005/>

A study of nine modern carnivores indicate that one in four adults had suffered tooth breakage and that half of these breakages were of the canine teeth. The highest frequency of breakage occurred in the spotted hyena, which is known to consume all of its prey including the bone. The least breakage occurred in the [[African wild dog]]. The gray wolf ranked between these two.<ref name=valkenburgh1993a/><ref name=valkenburgh1988/> The eating of bone increases the risk of accidental fracture due to the relatively high, unpredictable stresses that it creates. The most commonly broken teeth are the canines, followed by the premolars, carnassial molars, and incisors. Canines are the teeth most likely to break because of their shape and function, which subjects them to bending stresses that are unpredictable in direction and magnitude.<ref name=valkenburgh1988/> The risk of tooth fracture is also higher when taking and consuming large prey.<ref name=valkenburgh1988/><ref name=desantis2015/>
 
In comparison to extant gray wolves, the extinct [[Beringian wolf|Beringian wolves]] included many more individuals with moderately to heavily worn teeth and with a significantly greater number of broken teeth. The frequencies of fracture ranged from a minimum of 2% found in the [[Northern Rocky Mountain wolf]] ''(Canis lupus irremotus)'' up to a maximum of 11% found in Beringian wolves. The distribution of fractures across the tooth row also differs, with Beringian wolves having much higher frequencies of fracture for incisors, carnassials, and molars. A similar pattern was observed in spotted hyenas, suggesting that increased incisor and carnassial fracture reflects habitual bone consumption because bones are gnawed with the incisors and then cracked with the carnassials and molars.<ref name=leonard2007/>

== Coyotes, jackals, and wolves ==
The [[gray wolf]] (''C. lupus''), the [[Ethiopian wolf]] (''C. simensis''), [[eastern wolf]] (''C. lycaon''), and the [[African golden wolf]] (''C. lupaster'') are four of the many ''Canis'' species referred to as "wolves".<ref>{{Cite web |title=Wolf - Red, Eastern & Ethiopian Wolves, Extinct Falkland Islands & Dire Wolves {{!}} Britannica |url=https://www.britannica.com/animal/wolf/Other-wolves |access-date=2024-02-24 |website=www.britannica.com |language=en}}</ref> Species that are too small to attract the word "wolf" are called [[coyote]]s in the [[Americas]] and [[jackal]]s elsewhere.<ref>Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed February 24, 2024 at <nowiki>https://animaldiversity.org/accounts/Canis_latrans/</nowiki></ref> Although these may not be more closely related to each other than they are to ''C. lupus'', they are, as fellow ''Canis'' species, more closely related to wolves and domestic dogs than they are to [[fox]]es, [[maned wolf|maned wolves]], or other canids which do not belong to the genus ''Canis''. The word "jackal" is applied to the golden jackal (''C. aureus''), found across southwestern and south-central Asia, and [[the Balkans]] in Europe.<ref>{{Cite web |title=Canis aureus, Golden jackal |url=https://www.thainationalparks.com/species/golden-jackal |access-date=2024-02-24 |website=Thai National Parks |language=en }}</ref>

==African migration==
The first record of ''Canis'' on the African continent is ''Canis sp. A'' from South Turkwel, Kenya, dated 3.58–3.2&nbsp;million years ago.<ref>{{cite journal|doi=10.1111/j.1096-3642.2005.00165.x|title=Plio-Pleistocene Carnivora of eastern Africa: Species richness and turnover patterns|journal=Zoological Journal of the Linnean Society|volume=144|issue=2|pages=121|year=2005|last1=Werdelin|first1=Lars|last2=Lewis|first2=Margaret E|doi-access=free}}</ref> 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 colonised Africa from Eurasia at least 5 times throughout the [[Pliocene]] and [[Pleistocene]], which is consistent with fossil evidence suggesting that much of the African canid fauna diversity resulted from the immigration of Eurasian ancestors, likely coincident with [[Quaternary glaciation|Plio-Pleistocene climatic oscillations]] between arid and humid conditions.<ref>{{cite journal|doi=10.1016/j.cub.2015.06.060|pmid=26234211|title=Genome-wide Evidence Reveals that African and Eurasian Golden Jackals Are Distinct Species|journal=Current Biology|volume=25|issue=16|pages=2158–65|year=2015|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.|doi-access=free}}</ref>{{rp|S1}} In 2017, the fossil remains of a new ''Canis'' species, named ''Canis othmanii'', was discovered among remains found at Wadi Sarrat, Tunisia, from deposits that date 700,000 years ago. This canine shows a morphology more closely associated with canids from Eurasia instead of Africa.<ref>{{cite journal|doi=10.1016/j.crpv.2017.05.004|title=''Canis othmanii'' sp. nov. (Carnivora, Canidae) from the early Middle Pleistocene site of Wadi Sarrat (Tunisia)|journal=Comptes Rendus Palevol|volume=16|issue=7|pages=774|year=2017|last1=Amri|first1=Lamjed|last2=Bartolini Lucenti|first2=Saverio|last3=Mtimet|first3=Moncef Saïd|last4=Karoui-Yaakoub|first4=Narjess|last5=Ros-Montoya|first5=Sergio|last6=Espigares|first6=Maria-Patrocinio|last7=Boughdiri|first7=Mabrouk|last8=Bel Haj Ali|first8=Nebiha|last9=Martínez-Navarro|first9=Bienvenido|doi-access=}}</ref>

== Gallery ==
<gallery mode="packed">
File:Canis lupus signatus - 01.jpg|[[Gray wolf]] (''Canis lupus'')
File:Washtenaw County's last wolf (1907).jpg|[[Eastern wolf]] (''Canis lycaon'') (includes ''latrans'' admixture)
File:Red wolf (4531335218).jpg|[[Red wolf]] (''Canis rufus'') (includes ''latrans'' admixture)
File:USMC-11557.jpg|[[Coyote]] (''Canis latrans'')
File:Golden wolf small.jpg|[[African wolf]] (''Canis lupaster'')
File:Flickr - Rainbirder - Golden Jackal.jpg|[[Golden jackal]] (''Canis aureus'')
File:Canis simensis.jpg|[[Ethiopian wolf]] (''Canis simensis'')
File:Tibetan Wolf By Stanzin (Stakpa).jpg|[[Himalayan wolf]] (''Canis lupus chanco'')
File:Indian Wolf Male.jpg|[[Indian wolf]] (''Canis lupus pallipes'')
File:20110425 German Shepherd Dog 8505.jpg|[[Domestic dog]] (''Canis familiaris'')
</gallery>

== See also ==
* [[List of canids]]

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==External links==
{{Wikispecies|Canis}}
{{Wiktionary|Canis}}

{{Carnivora|Ca.}}
{{Canidae extinct nav|W.}}
{{Taxonbar|from=Q149892}}
{{Authority control}}

[[Category:Canis| ]]
[[Category:Canines]]
[[Category:Carnivoran genera]]
[[Category:Extant Tortonian first appearances]]
[[Category:Taxa named by Carl Linnaeus]]