Canis

Canis is a genus of the Caninae containing multiple extant species,

such as wolves, dogs, coyotes and jackals. Species of this genus Canis
are distinguished by their moderate to large size, their massive, Temporal range: Miocene–present
well-developed skulls and dentition, long legs, and comparatively (6 million years ago)[1]
short ears and tails.[6]

Contents
Taxonomy
Evolution
Dentition and biteforce
Behavior
Description and sexual dimorphism
Mating behaviour
Tooth breakage
Coyotes, jackals, and wolves
African migration
Gallery
Gray wolf (top), coyote and African
See also
golden wolf (top middle), Ethiopian
References
wolf and Eurasian golden jackal
External links
(bottom middle), black-backed jackal
and side-striped jackal (bottom)
Taxonomy Scientific classification
Kingdom: Animalia
The genus Canis (Carl Linnaeus, 1758) was published in the 10th
edition of Systema Naturae[2] and included the dog-like Phylum: Chordata
carnivores: the domestic dog, wolves, coyotes and jackals. All
Class: Mammalia
species within Canis are phylogenetically closely related with 78
chromosomes and can potentially interbreed.[7] In 1926, the Order: Carnivora
International Commission on Zoological Nomenclature (ICZN) in
Family: Canidae
Opinion 91 included Genus Canis on its Official Lists and Indexes
of Names in Zoology.[8] In 1955, the ICZN's Direction 22 added Subfamily: Caninae
Canis familiaris as the type specimen for genus Canis to the
Tribe: Canini
official list.[3]
Subtribe: Canina

Canis is primitive relative to Cuon, Lycaon, and Genus: Canis

Xenocyon in its relatively larger canines and lack of Linnaeus, 1758[2]
such dental adaptations for hypercarnivory as m1–m2
Type species
metaconid and entoconid small or absent; M1–M2
 hypocone small; M1–M2 lingual cingulum weak; M2 Canis familiaris (dog)[3][4][5]
and m2 small, may be single-rooted; m3 small or
absent; and wide palate. Extant species

— Richard H. Tedford[4] Canis adustus

Canis anthus
The cladogram below is based on the DNA phylogeny of
Lindblad-Toh et al. (2005),[9] modified to incorporate recent Canis aureus
findings on Canis species,[10] Canis latrans
Canis lupus
Canis lupus familiaris (domestic dog) Canis mesomelas
Canis simensis

Canis lupus (gray wolf)

Canis latrans (coyote)

Canis anthus (African golden wolf)

Canis simensis (Ethiopian wolf)

Canina

Canis aureus (Golden jackal)

Cuon alpinus (dhole)

Lycaon pictus (African wild dog)

Canis adustus (side-striped jackal)

Canis mesomelas (black-backed jackal)

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.[11]
Evolution

The fossil record shows that feliforms and caniforms emerged within the clade Carnivoramorpha 43 million
YBP.[12] The caniforms included the fox-like genus Leptocyon whose various species existed from 24
million YBP before branching 11.9 million YBP into Vulpes (foxes) and Canini (canines). The jackal-sized
Eucyon existed in North America from 10 million YBP and by the Early Pliocene about 6-5 million YBP the
coyote-like Eucyon davisi[13] invaded Eurasia. In North America it gave rise to early Canis which first
appeared in the Miocene (6 million YBP) in south-western USA and Mexico. By 5 million YBP the larger
Canis lepophagus appeared in the same region.[1]:p58

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. The position of the canids would change with
the arrival of Canis to become a dominant predator across the Skulls of dire wolf (C. dirus), gray
Holarctic. The wolf-sized C. chihliensis appeared in northern China wolf (C. lupus), eastern wolf (C.
in the Mid-Pliocene around 4-3 million YBP. This was followed by lycaon), red wolf (C. rufus), coyote
an explosion of Canis evolution across Eurasia in the Early (C. latrans), African golden wolf (C.
anthus), golden jackal (C. aureus)
Pleistocene around 1.8 million YBP in what is commonly referred to
and black-backed jackal (C.
as the wolf event. It is associated with the formation of the mammoth
mesomelas)
steppe and continental glaciation. Canis spread to Europe in the
forms of C. arnensis, C. etruscus, and C. falconeri.[1]:p148

Xenocyon (strange wolf) is an extinct subgenus of Canis.[14] 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 Canis (Xenocyon)
lycaonoides.[15] The hypercarnivore Xenocyon gave rise to the modern dhole and the African wild
dog.[1]:p149

See further: Evolution of the canids

Dentition and biteforce

Bite force adjusted for body weight in Newtons per
kilogram[16]
Canid Carnassial Canine
Wolf 131.6 127.3
Dhole 130.7 132.0
African wild dog 127.7 131.1
Greenland dog and Dingo 117.4 114.3
Coyote 107.2 98.9
Side-striped jackal 93.0 87.5
Golden jackal 89.6 87.7 Diagram of a wolf skull with key features labelled
Black-backed jackal 80.6 78.3

Dentition relates to the arrangement of teeth in the mouth, with the dental notation for the upper-jaw teeth
using the upper-case letters I to denote incisors, C for canines, P for premolars, and M for molars, and the
lower-case letters i, c, p and m to denote the mandible teeth. Teeth are numbered using one side of the mouth
and from the front of the mouth to the back. In carnivores, the upper
premolar P4 and the lower molar m1 form the carnassials that are
used together in a scissor-like action to shear the muscle and tendon
of prey.[1]: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
Eurasian wolf skull
carnassial the trigonid is used for slicing and the talonid is used for
grinding. The ratio between the trigonid and the talonid indicates a
carnivore's dietary habits, with a larger trigonid indicating a
hypercarnivore and a larger talonid indicating a more omnivorous diet.[17][18] Because of its low variability,
the length of the lower carnassial is used to provide an estimate of a carnivore's body size.[17]

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 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.[19]

Behavior

Description and sexual dimorphism

There is little variance among male and female canids. Canids tend to live as monogamous pairs. Wolves,
dholes, coyotes, and jackals live in groups that include breeding pairs 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.[20] Social
mammal predators prey on herbivores with a body mass similar to that of the combined mass of the predator
pack.[21][22] The gray wolf specializes in preying on the vulnerable individuals of large prey,[23] and a pack
of timber wolves can bring down a 500 kg (1,100 lb) moose.[24][25]

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.[26] 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. Wolves are typically monogamous and form 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 monopolize 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.[26]
Another study on free-ranging dogs 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.[27] They
found that females in heat searched out 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.[27]

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 intersexual parental investment.[28] 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.[29] 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.[30] 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 gray wolves relates to their mating system.

Tooth breakage

Tooth breakage is a frequent result of carnivores' feeding behaviour.[31]

Carnivores include both pack hunters 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 – a pack hunter or a solitary hunter – and
Dentition of a wolf showing
even how it consumed its prey. The mandibles of canids are buttressed
functions of the teeth.
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. 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.[32]

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.[31][33] 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.[33] The risk of tooth fracture is also higher when taking and
consuming large prey.[33][34]

In comparison to extant gray wolves, the extinct 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.[35]

Coyotes, jackals, and wolves

Wolves, dogs, and dingoes are all subspecies of Canis lupus.[5] The gray wolf (C. lupus), the Ethiopian wolf
(C. simensis), and the African golden wolf (C. anthus) are three of the many Canis species referred to as
"wolves"; however, all of the others are now extinct and little is known about them by the general public.
One of these, the extinct dire wolf (C. dirus), has gained fame from the thousands of specimens found and
displayed at the Rancho La Brea Tar Pits in Los Angeles, California.

Canis species that are too small to attract the word "wolf" are called coyotes in the Americas and jackals
elsewhere. Although these may not be more closely related to each other than they are to C. lupus, they are,
as fellow Canis species, all more closely related to wolves and domestic dogs than they are to foxes, maned
wolves, or other canids which do not belong to the genus Canis. The word "jackal" is applied to three
distinct species of this group: the side-striped (C. adustus) and black-backed (C. mesomelas) jackals, found
in sub-Saharan Africa, and the Eurasian golden jackal (C. aureus), found across southwestern and south-
central Asia, and the Balkans.

While North America has only one small-sized species, the coyote (C. latrans), it has become very
widespread, moving into areas once occupied by wolves. They can be found across much of mainland
Canada, in every state of the contiguous United States, all of Mexico except the Yucatán Peninsula, and the
Pacific and central areas of Central America, ranging as far as western Panama.

African migration
The first record of genus Canis on the African continent is Canis sp. A from South Turkwel, Kenya dated
3.58–3.2 million years ago.[36] 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 African canid fauna diversity resulted from the immigration of Eurasian
ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid
conditions.[37]: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 rather than Africa.[38]

Gallery
Gray wolf (Canis Eastern wolf (Canis Red wolf (Canis Coyote (Canis
lupus) (includes dog lycaon) (often rufus) (includes latrans)
and dingo). includes latrans latrans admixture)
admixture)

Dire wolf (Canis African golden wolf Eurasian golden Ethiopian wolf
dirus) (extinct) (Canis anthus) jackal (Canis (Canis simensis)
aureus)

Black-backed jackal Side-striped jackal

(Canis mesomelas) (Canis adustus)

See also
List of Canis species
List of canids

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