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Canisdogs, jackals, and wolves

By Lydia Oliver

Diversity

Canis is a diverse genera encompassing 7 species of canids which include jackals, wolves, coyotes, and many subspecies such as the domestic dog (Hailer and Leonard, 2008). Within the Order Carnivora, this group can be distinguished from others by their relatively large size, elongated rostrum with a well developed skull, including dentition categorized by large canine teeth and a pronounced carnassial pair (Loveridge and Macdonald, 2003). This predatory genus feeds on other animals - from small rodents to large herbivores. Given the opportunity they also can consume plant matter making them facultative carnivores (Wilson et al., 2012). Canids are top predators within the ecosystems they occupy - their diversity enables them to dominate desert, forest, and mountain habits (Wynne and Udell, 2010). (Hailer and Leonard, 2008; Loveridge and Macdonald, 2003; Wilson, et al., 2012; Wynne and Udell, 2010)

Geographic Range

Canis can be found dispersed throughout the Northern and Southern hemispheres (Beschta and Ripple, 2009). Canid species are native to the Neoartic, Paleartic, and Ethiopian Regions. Subspecies of Canis have been introduced to Neotropical, Oriental, and Australian regions (Gopalakrishnan et al., 2018). (Beschta and Ripple, 2009; Gopalakrishnan, et al., 2018)

Habitat

Canis occupy many habitats in the Old World and New World. Habitat variety include: dry open grasslands in Southern latitudes, dense boreal forests in higher Northern Latitudes, and even urban environments (Hailer and Leonard, 2008 & Humer, Heltai, and Murariu, 2012). The elevation distribution for canids range from sea level up to roughly 3,700 meters above sea level (Malcolm, 2004). Most commonly occupied habitats by canids include: forests, shrublands, inland wetlands, and rocky areas (Loveridge and Macdonald, 2003 & Malcolm, 2004). (Hailer and Leonard, 2008; Humer, et al., 2012; Loveridge and Macdonald, 2003; Malcolm, 2004)

Systematic and Taxonomic History

Canis encompasses "dog like" canids (Wilson et al., 2012). They are distinguished by their moderate to large size, highly developed skulls and dentition and long legs compared to body size (Meiri et al., 2007). Their unique dentition includes enlarged canine teeth plus a pair of molars used for sheering called a carnassial pair (Tanis, DeSantis, and Terry, 2018). Canis includes wolves, coyotes, and jackals (Phillips, 2018). Previously the Canis included foxes but they were removed and separated into their own Vulpes. The closest relatives to Canis are the foxes Vulpes (Van Valkenburgh et al., 2014). Other families that are similar to the Canis are the Mustelidae (weasels), Mephitidae (skunks), and Ursidae (bears) (Van Valkenburgh et al., 2014). (Meiri, et al., 2007; Phillips, 2018; Tanis, et al., 2018; Van Valkenburgh, et al., 2014; Wilson, et al., 2012)

  • Synapomorphies
    • Elongated Rostrum
    • Large Canine Teeth

Physical Description

Canis spcies can range in weight from 10 kg to 70 kgs (Malcolm, 2004). Coyotes Canis latrans and jackals Canis aureus & Canis adustus are in the smaller size range. The Ethiopian wolf Canis simensis, red wolf Canis rufus and the grey wolves Canis lupus are all significantly larger in height and weight with males typically over 60 kgs (Marafina Vieira Porto, Maestri, and Da Silva Duarte, 2019).

Male and female canids are all similar in looks, with males slightly larger in body mass than females (Wynne and Udell, 2010). Coat patterns and colors vary among species but common coat colors include: brown, black, grey, white, and red (Malcolm, 2004 & Meiri et al., 2007). Seasonal coat changes occur in canids that reside at high latitudes such as C. lupus and C. latrans.

Common features of Canids include an elongated rostrum, with large canine teeth, and a broad gait which makes covering long distances in search of prey more efficient (Van Valkenburgh, 1988). (Malcolm, 2004; Marafina Vieira Porto, et al., 2019; Meiri, et al., 2007; Van Valkenburgh, 1988; Wynne and Udell, 2010)

  • Sexual Dimorphism
  • sexes alike
  • male larger

Reproduction

The Canis species are unique among mammals because of the high incidence of monogamy (Feinstein et al., 2013). Most canids live in packs consisting of less than 10 individuals, where the dominant male and female pairs are the only ones to breed. Mates are first selected around the age of sexual maturity which typically occurs at 2-3 years of age (Malcolm, 2004). Male and female Canids both leave their natal pack in search of non related mates and to join a new pack (Wilson et al., 2012). (Feinstein, et al., 2013; Malcolm, 2004; Wilson, et al., 2012)

Mating occurs in spring and summer, generally from March to September. Canids in lower latitudes breed later in the summer months and higher latitudes breed in late spring. The gestation period of Canids is around 60 days and a typical litter size ranges from 2-6 pups. The young are born altricial within a den and develop motile skills within the first few weeks. After a few months, the pups slowly start to explore outside of the den but are not fully weaned from the mother until 8 weeks of age. Around the age of sexual maturity (2-3 years) Canids will either stay in their natal pack or leave to join another one in the hopes of finding a mate. (Feinstein, et al., 2013; Hailer and Leonard, 2008; Humer, et al., 2012)

  • Key Reproductive Features
  • gonochoric/gonochoristic/dioecious (sexes separate)
  • sexual

Maternal and paternal investment within Canis species is high (Malcolm, 2004). C. simensis and C. rufus exhibit polygyandry with multiple breeding pairs. Young are born altricial and need care for the first few months of life (Feinstein et al., 2013). Canids exhibit cooperative breeding where other pack members help to care for young (Marafina Vieira Porto, Maestri, and Da Silva Duarte, 2019).

Pups are dependent on the mother for milk until the age of 8 weeks, but other members of the pack begin to regurgitate food starting at 4 weeks of age (Feinstein et al., 2013). Males care for young by regurgitating food for the pups as well as bringing back food to the den for the alpha female to consume while she is still nursing (Marafina Vieira Porto, Maestri, and Da Silva Duarte, 2019). Pups are fully weaned by the age of 8 weeks. Pups are dependent on the pack for the first year of life to bring food either in the form of regurgitated matter or a carcass to learn necessary hunting skills (Malcolm, 2004). (Feinstein, et al., 2013; Malcolm, 2004; Marafina Vieira Porto, et al., 2019)

  • Parental Investment
  • altricial
  • male parental care
  • female parental care
  • pre-weaning/fledging
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • post-independence association with parents
  • extended period of juvenile learning

Lifespan/Longevity

Canis species are top predators in the ecosystems they inhabit and can live 6-8 years in the wild (Beschta and Ripple, 2009). Canids are sometimes killed by other canids in disputes over territory or mates (Wilson et al., 2012).

Living in groups make life easier for canids because they have protection within the group, can take down larger prey, and can hold territory without expending too much energy (Wynne and Udell, 2010). Most mortality occurs within the first few years of life when young are more susceptible to disease or starvation (Mladenoff et al., 1995). Natural mortality of adults can be the result of injuries from hunting or defending territory (Marino and Sillero-Zubiri, 2011). Most mortality in canids is caused by humans through wildlife conflicts over livestock death or agricultural crop raids (Malcolm, 2004). (Beschta and Ripple, 2009; Malcolm, 2004; Marino and Sillero-Zubiri, 2011; Mladenoff, et al., 1995; Wilson, et al., 2012; Wynne and Udell, 2010)

Behavior

Canis are highly social animals, living in groups of 3 to 10 individuals with a strict social hierarchy (Meiri et al., 2007). Living in packs allows easier acquisition of resources including land and food without having to over exert themselves (Wynne and Udell, 2010). A pack will defend a broad territory and move throughout that space. Top ranking males and females are the only individuals allowed to breed, and often times once the young hit sexual maturity, they leave their natal pack in search of new canids with which to form packs (Malcolm, 2004). (Malcolm, 2004; Meiri, et al., 2007; Wynne and Udell, 2010)

Communication and Perception

Canis have an acute sense of smell, well developed eyesight, and a broad range of vocal cues (Malcolm, 2004). These sensory adaptations help them to communicate through scent marking, body language, and vocalizations (Atkins and Dillion, 1971). Within close range, many canids use a variety of piloerection, tail flicks, ear positions, lip, jowl movements, and entire body movements to send cues for others to pick up information (Marafina Vieira Porto, Maestri, and Da Silva Duarte, 2019). Canids are tactile beings who utilize physical contact to communicate positive and negative feelings with other members of their pack (Atkins and Dillion, 1971).

Canids use their strong sense of smell the most - from finding food sources, identifying scent markings, and even to determine a new competitor, predator, or pack mate in the area (Meiri et al., 2007). At a distance, canids rely on a variety of vocalizations to communicate to other canids, and most vocalizations are species specific. Canis lupus howls to communicate. Canis latrans, Canis adustus, Canis mesomelas, and Canis aureus produce a variety of yips and small howls in different tones and pitches to communicate (Malcolm, 2004). (Atkins and Dillion, 1971; Malcolm, 2004; Marafina Vieira Porto, et al., 2019; Meiri, et al., 2007)

Food Habits

Canis are falcultative carnivores (Kupczik and Stynder, 2012). Feeding mainly on animal flesh, they supplement their diet with plant matter when available. Canids have a well developed pair of carnassials, designed for shearing meat and crushing bone (Kupczik and Stynder, 2012). Apex predator canids feed on mammals including elk Cervis canadensis, moose Alces alces, and deer Family Cervidae (Malcolm, 2004). Mesopredator canids feed on smaller mammalian prey including mice Peromyscus and rabbits Order Lagomorpha. Both apex and mesolevel canids supplement their diets by eating berries, fish, and some plant matter (Malcolm, 2004). Canids are hunters and scavengers, engaging in active pursuit of prey plus eating already deceased animals (Kupczik and Stynder, 2012). Weak or young prey are targeted during the hunt, when members of the pack flush prey out into the open and rush in to bite any possible part of the prey with the goal of taking down the animal. Once down, prey is torn into and canids will then gorge themselves on the carcass (Tanis, DeSantis, and Terry, 2018). (Kupczik and Stynder, 2012; Malcolm, 2004; Tanis, et al., 2018)

Predation

Canis are within top trophic levels for every ecosystem they occupy and have few natural predators outside of other canids (Van Valkenburgh, 1988). Larger canids will prey on smaller canids in the same habitat. Smaller canids will move locations to avoid conflicts with larger canids (Marafina Viera Porto, Masestri, & Da Silva Duarte, 2019). The biggest threat to canids are humans who view them as pests and seek lethal methods to mitigate and reduce conflict (Malcolm, 2004). (Malcolm, 2004; Marafina Vieira Porto, et al., 2019; Van Valkenburgh, 1988)

  • Known Predators
    • Humans

Ecosystem Roles

Canis are top and meso-level predators within the ecosystems they inhabit (Van Valkenburgh, 1988). They exercise top down control on prey populations and lower level predators in the trophic pyramid (Malcolm, 2004). (Malcolm, 2004; Van Valkenburgh, 1988)

Economic Importance for Humans: Positive

There is some ecotourism revolving around Canis and as a result positive economic potential and benefits (Malcolm, 2004). Canis species are also often sought out in the illegal pet trade as well as in the wild for their pelts (Phillips, 2018). (Malcolm, 2004; Phillips, 2018)

Economic Importance for Humans: Negative

Canis were viewed mainly in a positive light until the spread of agriculture put humans and canids in close proximity (Malcolm, 2004). Domestic livestock such as chickens, goats, and cows provide easy meals for opportunistic canids to eat (Mech, 1995). For some canids, like coyotes C. latrans, cultivated fruit are a favorite crop to eat. Economic loss from livestock and crop destruction has caused people to view them as pests (Mech, 1995). (Malcolm, 2004; Mech, 1995)

  • Negative Impacts
  • crop pest

Conservation Status

Majority of Canis species are growing in population numbers (Malcolm, 2004). But one species is endangered and another is critically endangered.

Ethiopian wolf C. simensis is labeled as endangered and decreasing in population, there are 197 mature adults in the wild (Marino and Sillero-Zubiri, 2011). The majority of Ethiopian wolves live within the Bale Mountains in Ethiopia (Malcolm, 2004). Living at high population density for large social predators makes it easier for diseases such as Canine Distemper Virus to effect the population, which is a large concern for the dwindling population (Marino and Sillero-Zubiri, 2011).

Red wolf C. rufus is labeled as critically endangered with populations decreasing (Phillips, 2018). There are thought to be 20-30 adults in the wild remaining from the reintroduction efforts in the late 1980s. Major issues concerning the Red wolf are hybridization with coyotes that also live within the area, habitat destruction, and human conflict issues within the South Eastern United States (Phillips, 2018). (Malcolm, 2004; Marino and Sillero-Zubiri, 2011; Phillips, 2018)

  • IUCN Red List [Link]
    Not Evaluated

Other Comments

Canis means "dog" in Latin, "canine tooth" is also derived from this term due to the long tooth that all canids possess (Van Valkenburgh et al., 2014). Companion dogs C. lupus familiaris have played a significant role in human history for thousands of years, and our bond with companion dogs have led to a fascination with wild canids (Malcolm, 2004). (Malcolm, 2004; Van Valkenburgh, et al., 2014)

Contributors

Lydia Oliver (author), Colorado State University, Genevieve Barnett (editor), Colorado State University.

Glossary

Australian

Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.

World Map

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

cooperative breeder

helpers provide assistance in raising young that are not their own

crepuscular

active at dawn and dusk

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

duets

to jointly display, usually with sounds in a highly coordinated fashion, at the same time as one other individual of the same species, often a mate

ecotourism

humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

introduced

referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.

male parental care

parental care is carried out by males

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nomadic

generally wanders from place to place, usually within a well-defined range.

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

piscivore

an animal that mainly eats fish

scavenger

an animal that mainly eats dead animals

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

scrub forest

scrub forests develop in areas that experience dry seasons.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

tundra

A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.

urban

living in cities and large towns, landscapes dominated by human structures and activity.

visual

uses sight to communicate

References

Atkins, D., L. Dillion. 1971. Evolution of the Cerebellum in the Genus Canis. Journal of Mammalogy, 52: 96-107.

Beschta, R., W. Ripple. 2009. Large Predators and Trophic Cascades in Terrestrial Ecosystems of the Western United States. Biological Conservation, 142: 2401-2414.

Feinstein, M., B. Smith, R. Coppinger, K. Lord. 2013. Variation in Reproductive Traits of Members of the Genus Canis with Special Attention to the Domestic Dog (Canis familiaris). Journal of Behavioural Processes, 92: 131-142.

Gopalakrishnan, S., M. Sinding, J. Ramos-Madrigal, O. Wiig, A. Hansen, M. Gilbert. 2018. Interspecific Gene Flow Shaped the Evolution of the Genus Canis. Current Biology, 28, 21: 3441-3449.

Gronau, I., R. Schweizer, D. Ortega-Del Vecchyo, E. Han, A. Freedman, A. Boyko. 2014. Genome Sequencing Highlights the Dynamic Early History of Dogs. PLOS Genetics, 10: 1.

Hailer, F., J. Leonard. 2008. Hybridization Among Three Native North American Canis Species in a Region of Natural Sympatry. PLOS One, 3: 10.

Humer, A., M. Heltai, D. Murariu. 2012. Current Status and Distribution of Golden Jackals, Canis aureus in Europe. Mammalogy Review, 42: 1-11.

Kupczik, K., D. Stynder. 2012. Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora). Biological Journal of the Linnean Society, 105: 456-471.

Loveridge, A., D. Macdonald. 2003. Niche Separation in Sympatric Jackals (Canis mesomelas and Canis adustus). The Zoological Society of London, 259: 143-153.

Malcolm, J. 2004. Dogs, Wolves, Coyotes, Jackals, and Foxes (Canidae) Citation metadata. Pp. 254 in M Hutchins, A Evans, J Jackson, D Kleiman, J Murphy, D Thoney, eds. Grzimek's Animal Life Encyclopedia, Vol. 14, 2 Edition. Detroit, MI: Gale.

Marafina Vieira Porto, L., R. Maestri, L. Da Silva Duarte. 2019. Evolutionary Relationships Among Life-History Traits in Caninae (Mammalia:Carnivora). Biological Journal of the Linnean Society, 128: 311-322.

Marino, J., C. Sillero-Zubiri. 2011. "IUCN Redlist" (On-line). Accessed March 08, 2020 at https://www.iucnredlist.org/species/3748/10051312#population.

Mech, D. 1995. The Challenge and Opportunity of Recovering Wolf Population. Society for Conservation Biology, 9,2: 270-278.

Meiri, S., T. Barrackough, J. Gittleman, J. Davies. 2007. Species Coexistence and Character Divergence Across Carnivores. Ecology, 10: 146-152.

Mladenoff, D., T. Sickley, R. Haight, A. Wydeven. 1995. A Regional Landscape Analysis and Prediction of Favorable Gray Wolf Habitat in the Northern Great Lakes Region. Society for Conservation Biology, 9,2: 279-294.

Phillips, M. 2018. "IUCN Redlist" (On-line). Accessed March 08, 2020 at https://www.iucnredlist.org/species/3747/119741683.

Tanis, B., L. DeSantis, R. Terry. 2018. Dental microwear textures across cheek teeth in canids: Implications for dietary studies of extant and extinct canids. Journal of Palaeogeography, Palaeoclimatology, Palaeoecology, 508: 129-138.

Van Valkenburgh, B. 1988. Trophic Diversity in Past and Present Guilds of Large Predatory Mammals. Journal of Paleobiology, 14/2: 155-173.

Van Valkenburgh, B., B. Pang, D. Bird, A. Curtis, K. Yee, C. Wysocki, B. Craven. 2014. Respiratory and Olfactory Turbinals in Feliform and Caniform Carnivorans: the Influence of Snout Length. The Anatomical Record, 10.1002: 297.

Wilson, P., C. Kluetsch, B. Patterson, B. White, L. Rutledge. 2012. Conservation Genomics in Perspective: A Holistic Approach to Understanding Canis Evolution in North America. Biological Conservation, 155: 186-192.

Wynne, C., M. Udell. 2010. Ontogeny and Phylogeny: Both are Essential to Human-Sensitive Behaviour in the Genus Canis. Animal Behaviour, 79,2: 9-14.

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