Lifespan, longevity, and ageing

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Maximum longevity: 21.8 years (captivity)
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Joao Pedro de Magalhaes
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de Magalhaes, J. P.
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Untitled

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Coyotes are one of the dominant terrestrial carnivores in North America, with humans and wolves being their greatest enemies.

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
author
Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Behavior

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Coyotes use auditory, visual, olfactory and tactile signals to communicate. They are the most vocal of all North American wild mammals, using 3 distinct calls (squeak, distress call and howl call) which consist of a quick series of yelps, followed by a falsetto howl. Howling may act to announce where territories are to other packs. Coyotes also howl when two or more members of a pack re-unite and to announce to each other their location. Their sight is less developed and is used primarily to note movement. They have acute hearing and sense of smell. They use stumps, posts, bushes or rocks as "scent posts" on which they urinate and defecate, possibly to mark territory. Coyotes are very good swimmers but poor climbers.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: choruses ; scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Conservation Status

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Coyotes are common and widespread because of their extraordinary adaptability.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: least concern

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Benefits

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Coyotes serves as hosts for a number of diseases, including rabies. They are considered a threat to poultry, livestock, and crops. Coyotes may also compete with hunters for deer, rabbits, and other game species.

Negative Impacts: injures humans (carries human disease); causes or carries domestic animal disease

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Benefits

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Coyotes help to control some agricultural pests, such as rodents. Coyote pelts are also still collected and sold in some areas.

Positive Impacts: body parts are source of valuable material; controls pest population

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Associations

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Coyotes help in keeping many small mammal populations in check, such as mice and rabbits. If populations of these small mammals were allowed to become too large it would result in habitat degradation

Mutualist Species:

  • American badgers (Taxidea taxus)
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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Trophic Strategy

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Coyotes are versatile in their eating habits. They are carnivorous; 90% of their diet is mammalian. They eat primarily small mammals, such as eastern cottontail rabbits, thirteen-lined ground squirrels, and white-footed mice. They occasionally eat birds, snakes, large insects and other large invertebrates. They prefer fresh meat, but they consume large amounts of carrion. Part of what makes coyotes so successful at living in so many different places is the fact that they will eat almost anything, including human trash and household pets in suburban areas. Plants eaten include leaves of balsam fir and white cedar, sasparilla, strawberry, and apple. Fruits and vegetables are a significant part of the diet of coyotes in the fall and winter months. Coyotes hunt animals in interesting ways. When on a "mousing" expedition, they slowly stalk through the grass and sniff out the mouse. Suddenly, with all four legs held stiffly together, the coyotes stiffen and pounce on the prey. Hunting deer, on the other hand, calls for teamwork. Coyotes may take turns pursuing the deer until it tires, or they may drive it towards a hidden member of the pack. Coyotes sometimes form "hunting partnerships" with badgers. Because coyotes aren't very effective at digging rodents out of their burrows, they chase the animals while they're above ground. Badgers do not run quickly, but are well-adapted to digging rodents out of burrows. When both hunt together they effectively leave no escape for prey in the area. The average distance covered in a night's hunting is 4 km.

Animal Foods: birds; mammals; reptiles; carrion ; insects; terrestrial non-insect arthropods; terrestrial worms

Plant Foods: leaves; fruit

Primary Diet: omnivore

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Distribution

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Coyotes are native to the Nearctic region. They are found throughout North and Central America. They range from Panama in the south, north through Mexico, the United States, and Canada. They occur as far north as Alaska and all but the northernmost portions of Canada.

Biogeographic Regions: nearctic (Native )

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Habitat

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Coyotes are extremely adaptable and use a wide range of habitats including forests, grasslands, deserts, and swamps. They are typically excluded from areas with wolves. Coyotes, because of their tolerance for human activities, also occur in suburban, agricultural, and urban settings.

Habitat Regions: temperate ; terrestrial

Terrestrial Biomes: taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; scrub forest ; mountains

Wetlands: swamp

Other Habitat Features: urban ; suburban ; agricultural ; riparian

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Life Expectancy

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Coyotes have been known to live a maximum of ten years in the wild and 18 years in captivity.

Range lifespan
Status: wild:
10 (high) years.

Range lifespan
Status: captivity:
18 (high) years.

Average lifespan
Status: captivity:
15.0 years.

Average lifespan
Status: wild:
14.5 years.

Average lifespan
Status: captivity:
21.8 years.

Average lifespan
Status: wild:
21.0 years.

Average lifespan
Status: wild:
14.5 years.

Average lifespan
Status: captivity:
18.0 years.

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Morphology

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Coloration of coyotes varies from grayish brown to a yellowish gray on the upper parts. The throat and belly are whitish. The forelegs, sides of head, muzzle and feet are reddish brown. The back has fulvous colored underfur and long, black-tipped guard hairs that produce a black dorsal stripe and a dark cross on the shoulder area. The tail, which is half the body length, is bottle shaped with a black tip. There is also a scent gland located on the dorsal base of the tail. There is one moult per year, which starts in May with light loss of hair and ends in July after profuse shedding. Coyotes are significantly smaller than gray wolves and much larger than foxes. Coyotes are distinguished from domesticated dogs by their pointed, erect ears and drooping tail, which they hold below their back when running. The eyes have a yellow iris and round pupil. The nose is black and usually less than one inch in diameter. The ears are large in relation to the head and the muzzle is long and slender. The feet are relatively small for the size of the body. The pes has four digits and the manus has five with a small first digit. Coyotes run on their toes (digitigrade). The dental formula is 3/3 1/1 4/4 2/3. The molars are structured for crushing and the canines are rather long and slender.

Range mass: 7 to 21 kg.

Range length: 75 to 100 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

Average basal metabolic rate: 19.423 W.

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Associations

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Coyotes are very secretive. Especially near human habitations they are active mostly early in the morning and late in the evening. Coyotes keep their young in or near the den while they are young so that the pups aren't killed by predators and competitors such as wolves and mountain lions.

Known Predators:

  • humans (Homo sapiens)
  • gray wolves (Canis lupus)
  • mountain lions (Puma concolor)
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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Reproduction

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Courtship lasts for approximately 2 to 3 months. Female coyotes are monoestrous and are in heat for 2 to 5 days between late January and late March. Mating occurs within these 3 months. Once the female chooses a partner, the mates may remain paired for a number of years, but not necessarily for life.

Mating System: monogamous

Spermatogenesis in males takes around 54 days and occurs between January and February depending on geographic location. Gestation lasts from 60 to 63 days. Litter size ranges from 1 to 19 pups; the average is 6. The pups weigh approximately 250 grams. The young are born blind, limp-eared and pug-nosed. After 10 days the eyes open, the pups weigh 600 grams and their ears begin to erect in true coyote fashion. Twenty-one to 28 days after birth, the young begin to emerge from the den and by 35 days they are fully weaned. They are fed regurgitated food by both parents. Male pups disperse from the dens between months 6 and 9, while females usually stay with the parents and form the basis of the pack. Adult size is reached between 9 and 12 months. Sexual maturity is reached by 12 months. Coyotes hybridize with domestic dogs and occasionally with gray wolves.

Breeding interval: Coyotes usually breed once each year.

Breeding season: Breeding occurs from January to March.

Range number of offspring: 1 to 19.

Average number of offspring: 5.7.

Range gestation period: 50 to 65 days.

Range weaning age: 35 to 49 days.

Range age at sexual or reproductive maturity (female): 9 to 10 months.

Range age at sexual or reproductive maturity (male): 9 to 10 months.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization ; viviparous

Average birth mass: 250 g.

Average number of offspring: 6.

Female coyotes gestate and nurse their young. Both male and female coyotes bring food to their young after they are weaned and protect their offspring. The young sometimes stay with the pack into adulthood and learn how to hunt during a learning period.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Male, Female); pre-independence (Provisioning: Male, Female, Protecting: Male, Female); post-independence association with parents; extended period of juvenile learning

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Tokar, E. 2001. "Canis latrans" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Canis_latrans.html
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Erik Tokar, University of Michigan-Ann Arbor
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Allison Poor, University of Michigan-Ann Arbor
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Tanya Dewey, Animal Diversity Web
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Associated Plant Communities

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More info for the term: tundra

Coyotes evolved in a plains environment and were historically most
numerous in western grasslands where large ungulate populations were
high. Coyotes flourished in the shortgrass-steppe, semiarid sagebrush
(Artemisia spp.)-grasslands, and deserts, and they ranged from deserts
and plains to alpine areas of adjacent mountains [58].

Today, range expansions indicate that coyotes can be successful in any
plant community from the tropics of Guatemala to the tundra of northern
Alaska [58]. Although they occur in most plant communities throughout
their range, coyotes do show some preferences. In the Intermountain
region, coyotes are closely associated with sagebrush communities.
Coyotes in eastern Nevada preferred black sagebrush (Artemisia nova)
flats to other habitats. These flats were areas of highest black-tailed
jackrabbit (Lepus californicus) densities [44]. In the Sierra Nevada,
California, coyotes inhabit almost every plant community and
successional stage. However, they prefer grass-forb and shrub-conifer
seedling-conifer sapling communities [63].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Common Names

provided by Fire Effects Information System Animals
coyote
brush wolf
prairie wolf
American jackal
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Conservation Status

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Information on state- and province-level protection status of animals in
the United States and Canada is available at NatureServe, although
recent changes in status may not be included.
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Cover Requirements

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More info for the terms: cover, forest

Coyotes commonly hunt in open to semiopen areas [12,18,51]. In
California coyotes used ecotones, fuelbreaks, roads, trails, and open
chaparral more than dense unbroken cover. In southern California where
chaparral is adjacent to unbroken areas, coyotes forage at night along
edges and return during the day to chaparral cover. The steep slopes
and heavy cover of most chaparral communities impede coyote movements
[51]. In Georgia, the proportion of open area in coyote home ranges was
significantly (P less than 0.04) greater than that generally available in the
area, and the proportion of forest was significantly (P less than 0.04) less [59].

Coyotes use cover for daytime resting and den sites. In Georgia, areas
with "sufficient" cover were used more for daytime rest sites, and early
successional and open areas were used more for nocturnal foraging. In
summer, some coyotes used corn fields for cover during the day [59].
Urban coyotes in Seattle, Washington, foraged in residential areas, but
only in areas that were immediately adjacent to forest cover. Forested
areas provided the majority of cover, including denning sites [51].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Distribution

provided by Fire Effects Information System Animals
Coyotes are found from Costa Rica to northern Alaska, and from coast to
coast in the United States and Canada. The highest densities occur in
the Great Plains states and in south-central United States. Coyotes are
absent from the barrens and Arctic islands of northern Canada, including
much of northern Quebec, northern Newfoundland, and Labrador. Coyotes
are uncommon where gray wolf populations are high in northeastern
Minnesota, northern Alaska, the Northwest Territories, Manitoba, and
Ontario. The distribution of coyotes in eastern North America has
expanded during this century. In some states such as Florida and
Georgia, coyotes have been introduced [4,12,43]. Today, all eastern
states and provinces have at least a small population of coyotes [64].
Distribution of the subspecies is listed below [61,66]:

Mexican coyote - Occurs in Oaxaca, San Luis Potosi, Pueblo, and
Veracrus, Mexico. Its range may extend into southern Nuevo Leon and
southern Tamaulipas, Mexico.

San Pedro Martir coyote - Occurs in northern Baja California and
southwestern California (mostly San Diego County).

southeastern coyote - Occurs in southeastern and extreme eastern Kansas,
Oklahoma, Texas, Missouri, and Arkansas.

Durango coyote - Occurs along the Pacific coast drainage of western
Mexico between about 22 degrees and 26 degrees north latitude, extreme
southern Sonora, extreme southwestern Chihuahua, western Durango,
western Zacatecas, and Sinaloa.

northern coyote - In Canada, northern coyotes occur in Yukon Territory,
the Northwest Territories, northern British Columbia, and northern
Alberta. In the United States, northern coyotes occur in most of Alaska
except the southeastern coastal section.

Tiburon Island coyote - Occurs on Tiburon Island off Baja California.

plains coyote - In Canada, plains coyotes occur in southeastern Alberta,
southern Saskatchewan, and the extreme southwestern corner of Manitoba.
In the United States, they occur in Montana, Wyoming, and Colorado east
of the Rocky Mountains, and the northeastern corner of New Mexico; North
Dakota except the northeastern quarter; northwestern Oklahoma, and the
northern Panhandle region of Texas.

mountain coyote - In Canada, mountain coyotes occur in southern British
Columbia and southeastern Alberta. In the United States, they occur in
Oregon and Washington east of the Cascade Range, northern California,
Idaho, western Montana, Wyoming, and Colorado (except the southeast
corner), northern and central Nevada, and northern and central Utah.

Mearns coyote - Occurs in southwestern Colorado, extreme southern Utah
and Nevada, southeastern California, northeastern Baja California,
Arizona, west of the Rio Grande in New Mexico, and Sonora and Chihuahua
in Mexico.

Lower Rio Grande coyote - Occurs in extreme southern Texas and northern
Tamaulipas, Mexico.

California valley coyote - Occurs in California west of the Sierra
Nevada, except in the northern part.

peninsula coyote - Occurs on the Baja California peninsula.

Texas plains coyote - Occurs in Texas, except for the northern panhandle
region, the eastern part, and the extreme southern tip. Texas plains
coyotes also occur in eastern New Mexico except for the northeastern
corner, and part of northeastern Mexico.

northeastern coyote - In Canada, northeastern coyotes occur in
north-central Saskatchewan, Manitoba (except the extreme southwestern
corner), southern Ontario, and extreme southern Quebec. In the United
States, northeastern coyotes occur along the eastern edge of North
Dakota and in Minnesota, Iowa, Missouri (north of the Missouri River),
Michigan, Wisconsin, Illinois (except the extreme southern portion), and
northern Indiana.

northwest coast coyote - Occurs west of the Cascade Range in Oregon and
Washington.

Colima coyote - Occurs along the southwestern Pacific slope of Jalisco,
Michoacan, and Guerrero, Mexico.
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Food Habits

provided by Fire Effects Information System Animals
More info for the term: fruit

Coyotes are opportunistic feeders and eat a variety of food [4,12,64].
About 90 percent of their diet consists of animal matter; however, they
also eat vegetable matter. Some common prey items include deer, elk,
sheep (Ovis spp.), rabbits and hares (Leporidae), various rodents
(Rodentia), ground-nesting birds, amphibians, lizards, snails, fish,
crustaceans, and insects. During winter, much of the diet is made up of
rabbits, hares, and the carrion of large ungulates. Small mammals,
especially voles and mice (Muridae), are important food items during
spring, summer, and fall [4,64]. Various berries are also eaten [4].

An extensive study of coyote food habits conducted in 17 western states
showed that major diet items were lagomorphs (33%), carrion (25%),
rodents (18%), and domestic livestock (13.5%) [56]. Coyote diets in
sagebrush habitat of northeastern Utah and south-central Idaho consisted
of about 75 percent black-tailed jackrabbits year-round [13]. In
northeastern California, meadow voles (Microtus pennsylvanicus) occurred
in about half of all coyote scats analyzed. Other important diet items
were mule deer (Odocoileus hemionus) and cattle, probably eaten as
carrion [33]. Mule deer were also important in coyote diets in two
areas of southern Utah. In central Wyoming, mule deer, pronghorn
(Antilocapra americana), white-tailed jackrabbits (Lepus townsendii),
and desert cottontails (Sylvilagus audubonii) were present in 63 percent
of coyote scats [58]. On Arizona cattle ranges, where the habitat was
primarily open grasslands, oak (Quercus spp.), juniper, and ponderosa
pine (Pinus ponderosa), coyote diets contained high percentages of plant
material. Juniper berries were particularly important, followed by
prickly pear (Opuntia spp.) fruit [47].
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bibliographic citation
Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat-related Fire Effects

provided by Fire Effects Information System Animals
More info for the terms: cover, density, fire exclusion, fire suppression, forest, wildfire

Fire may improve the foraging habitat and prey base of coyotes. In New
England, coyotes are commonly found in forest openings created by fire
or logging [18]. Fires that reduce vegetation height and create open
areas probably increase hunting efficiency by coyotes. Surface fires
often open substrates for quieter stalking and easier capture of prey
than can occur in closed forests [38]. Wirtz [68] noted increases in
consumption of birds and deer by coyotes after a chaparral fire in the
San Dimas Experimental Forest, California. Increased consumption was
presumably the result of increased vulnerability of prey with reduced
cover, but no change was noted in small mammal consumption.

Periodic fire helps to maintain habitat for many prey species of coyote.
Fires that create a mosaic of burned and unburned areas are probably the
most beneficial to many coyote prey species. Several studies indicate
that many small mammal populations increase rapidly subsequent to
burning in response to increased food availability. Fire often improves
hare and rabbit forage quality and quantity for two or more growing
seasons [38]. Hill [67] concluded that burning at intervals longer than
2 years would be less beneficial to rabbits and hares, but any fire is
believed better than fire exclusion. Along the coast of northern
California, black-tailed jackrabbits occurred at highest density in open
brush, moderate density on recent burn areas, and lowest density in
mature chaparral stands [68]. Wagle [65] reported that fire suppression
in grasslands is detrimental to populations of small bird and mammal
herbivores due to organic matter accumulation and reduced plant vigor.

The 1988 fires in Yellowstone National Park have probably benefited
coyotes. Fire in combination with drought likely increased available
carrion the fall and winter following the fire. Additionally, the fires
stimulated grass production, which should lead to an increase in small
mammal populations [45].

In California, coyotes are abundant in young chaparral (less than 20
years old) and are rare or absent in chaparral that has not been burned
for 20 years or more [51]. Quinn [51] observed more coyote sign during
the second and third years after a chamise (Adenostoma spp.) chaparral
wildfire in Riverside County than had been observed prior to burning.
Coyote numbers increased during the second and third years following a
chaparral fire in the Sierra Nevada foothills [39].
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bibliographic citation
Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Cover Types

provided by Fire Effects Information System Animals
More info on this topic.

This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the term: cover

Coyotes probably occur in all SAF cover types.
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

More info for the term: shrub

FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES17 Elm-ash-cottonwood
FRES18 Maple-beech-birch
FRES19 Aspen-birch
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES27 Redwood
FRES28 Western hardwoods
FRES29 Sagebrush
FRES30 Desert shrub
FRES31 Shinnery
FRES32 Texas savanna
FRES33 Southwestern shrubsteppe
FRES34 Chaparral-mountain shrub
FRES35 Pinyon-juniper
FRES36 Mountain grasslands
FRES37 Mountain meadows
FRES38 Plains grasslands
FRES39 Prairie
FRES40 Desert grasslands
FRES41 Wet grasslands
FRES42 Annual grasslands
FRES44 Alpine
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Plant Associations

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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

Coyotes probably occur in all Kuchler plant associations.
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Habitat: Rangeland Cover Types

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This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the term: cover

Coyotes probably occur in all SRM (rangeland) cover types.
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Management Considerations

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Coyotes are the principal predator of domestic sheep in the West [44].
Predation on sheep often occurs in the summer [64]. In 16 studies
reviewed by Sterner and Shumake [60], coyotes were responsible for 82
percent of all sheep losses due to predators. However, only a few
flocks typically showed sizeable losses [12]. Coyote predation is a
minor cause of most livestock losses. Most of the livestock consumed,
except sheep, is carrion [64].

Methods of coyote control have been described in the literature
[1,4,12,64]. The impact of predator control on coyote population
densities, behavior, and ecology are not well known. Coyote populations
are able to maintain themselves under considerable human-induced
mortality. Their means of survival include behavioral adaptations and
biological compensatory mechanisms such as increased rates of
reproduction, survival, and immigration. In most areas, coyote numbers
likely are controlled by competition for food and by social stress,
diseases, and parasites [1]. There is little evidence to support the
notion that coyote predation is a primary limiting factor on populations
of large ungulates [12].

Coyote population control efforts may affect the social organization and
activity patterns of coyotes. In areas where population control is not
practiced, most coyotes exist in relatively "large" groups, whereas
coyotes in areas where populations are controlled generally exist in
"smaller" groups. Coyotes have been reported as more active during the
day in uncontrolled [26,70] than in population-controlled areas [71]. Roy
and Dorrance [72] reported that coyotes avoided open areas near roads
during daylight hours in areas where they were hunted.

Coyotes often aid in the dispersal of seeds. Seeds of oneseed juniper
(Juniperus monosperma) and Indian manzanita (Arctostaphylos mewukka)
have been found in coyote scats [24,31].

Coyotes are inflicted with a wide variety of parasites and diseases
which are described by Gier and others [28].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Occurrence in North America

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AL
AK
AZ
AR
CA
CO
CT
DE
FL
GA

ID
IL
IN
IA
KS
KY
LA
ME
MD

MA
MI
MN
MS
MO
MT
NE
NV
NH
NJ

NM
NY
NC
ND
OH
OK
OR
PA
RI
SC

SD
TN
TX
UT
VT
VA
WA
WV
WI
WY

AB
BC
MB
NB
NF
NT
NS
ON
PE
PQ

SK
YT
MEXICO

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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Predators

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Mountain lions (Felis concolor) sometimes kill and eat coyotes [4].
Other predators of coyotes include humans, gray wolves, black bears
(Ursus americanus), and grizzly bears (Ursus arctos). Golden eagles
(Aquila chrysaetos) attack young coyotes [2].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Preferred Habitat

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More info for the terms: cover, grassland, shrubland

Coyotes occupy a broad range of habitats [4,12,64]. Almost any habitat
that supports prey populations also supports coyotes; however, some
preferences have been noted (refer to PLANT COMMUNITIES slot) [64].

Dens - Coyotes den in a wide variety of places, including brush-covered
slopes, steep banks, rock ledges, thickets, and hollow logs. Dens
previously used by other animals (e.g., American badgers [Taxidea
taxus]) are frequently used [12]. Dens are usually about 1 foot (0.3 m)
in diameter and from 5 to 25 feet (1.5-7.5 m) long [4]. They usually
have more than one entrance and many interconnecting tunnels. The same
den may be used from year to year. Den sharing occurs only rarely
[4,12]. Movement of pups from one den to another is very common. The
reason is unknown, but disturbance and possibly infestation by parasites
may be factors. Most moves are over relatively short distances;
however, moves over 2.5 miles (4 km) are not uncommon [12].

Home range and territory - A single home range may be inhabited by a
family of two or more generations, a mated pair, or a single adult.
Home ranges vary from an average of 2 square miles (5 sq km) in Texas
[1] to averages of 21 to 55 square miles (54-142 sq km) in Washington
[57]. Males tend to have larger home ranges than females. In
Minnesota, male home ranges averaged 16 square miles (42 sq km), whereas
those of females averaged 4 square miles (10 sq km). The home ranges of
males overlapped considerably, but those of females did not [4]. In
Arkansas, Gipson and Sealander [26] reported that male coyote home
ranges were 8 to 16 square miles (21-42 sq km) and female home ranges
were 3 to 4 square miles (8-10 sq km).

In southeastern Colorado, the home range size of coyotes varied with
habitat, which was correlated with prey abundance. Coyotes in canyon
woodlands and in hills dominated by pinyon-juniper (Pinus-Juniperus
spp.) woodlands interspersed with grassland and shrubland had the
smallest home ranges. Coyotes in pinyon-juniper-prairie had
intermediate-size home ranges, and coyotes in shortgrass prairie had the
largest home ranges. As the amount of pinyon-juniper increased, home
range size decreased, possibly because these areas had high small mammal
populations and provided cover for resting sites and dens. The
shortgrass prairie had the lowest relative abundance of small mammals in
the study area [25].

Group size and social behavior may also influence home range size.
Coyotes living in packs and defending ungulate carrion during winter may
have smaller home ranges than coyotes living in pairs or alone [12,64].
Typically, only pack members defend territories; pairs of coyotes and
solitary individuals do not [4,12].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Taxonomy

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The currently accepted scientific name for the coyote is Canis latrans
Say. It is in the family Canidae. Nineteen subspecies are currently
recognized, however; only 16 subspecies occur in Mexico, the United
States, and Canada [4,30]:

Canis latrans cagottis (Hamilton-Smith) (Mexican coyote)
Canis latrans clepticus Elliot (San Pedro Martir coyote)
Canis latrans frustror Woodhouse (southeastern coyote)
Canis latrans impavidus Allen (Durango coyote)
Canis latrans incolatus Hall (northern coyote)
Canis latrans jamesi Townsend (Tiburon Island coyote)
Canis latrans latrans (plains coyote)
Canis latrans lestes Merriam (mountain coyote)
Canis latrans mearnsi Merriam (Mearns coyote)
Canis latrans microdon Merriam (Lower Rio Grande coyote)
Canis latrans ochropus Eschscholtz (California valley coyote)
Canis latrans peninsulae Merriam (peninsula coyote)
Canis latrans texesis Bailey (Texas plains coyote)
Canis latrans thamnos Jackson (northeastern coyote)
Canis latrans umpquesis Jackson (northwest coast coyote)
Canis latrans vigilis Merriam (Colima coyote)

Fertile hybrids have been produced by matings of coyotes with feral dogs
(C. familiaris), red wolves (C. rufus), gray wolves (C. lupus), and red
foxes (Vulpes vulpes) [4,12]. Coyote-dog hybrids exhibit decreased
fecundity [12].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Timing of Major Life History Events

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More info for the terms: density, litter, monoestrous

Social organization - There is a considerable amount of variability in
coyote social organizations. In many areas, most coyotes are solitary
outside of the breeding season. In other areas, such as Jackson Hole,
Wyoming, and Jasper, Alberta, groups of coyotes are frequently observed.
Coyote social organization is influenced by prey size. In populations
where the major prey items throughout the year are small rodents,
coyotes tend to be solitary. In populations where large animals are
available (e.g., elk [Cervus elaphus], and deer [Odocoileus spp.]),
large groups of coyotes form [12].

Breeding season - Courtship may begin as early as 2 to 3 months before
coyotes attempt to mate. The female is monoestrous, having one period
of heat per year usually between January and March [4,62]. Estrus lasts
2 to 5 days. Some coyotes mate with the same individual from year to
year, but not necessarily for life [4]. In the Sierra Nevada, coyotes
mate from February to May, with peak breeding time in April and May [63].
Yearling females usually breed later in the season than older females
[12].

Age at first breeding - Both males and females are capable of breeding
as yearlings [4]. However, many coyotes do not breed until their second
year [63]. Generally, about 60 to 90 percent of adult females and 0 to
70 percent of female yearlings produce litters [12]. In years when food
is abundant, more females (especially yearlings) breed. In years when
rodent populations are high, as many as 75 percent of yearling females
may breed [4].

Gestation and litter size - Gestation lasts approximately 63 days. The
average litter size is 6, but may range from 3 to 15 [12,63]. Litter
size can be affected by population density and food availability.
Knowlton [36] reported average litter sizes of 4.3 at high coyote
densities and 6.9 at low coyote densities. In years of high rodent
density, mean litter size is generally higher than in years of low
rodent densities [12].

Development of young - Coyote young are born with their eyes closed.
They are cared for by the mother and sometimes siblings from a previous
year. The father and other males often provide food for the mother and
the young. Pups emerge from the den in 2 or 3 weeks. They begin to eat
solid food at about 3 weeks of age and are weaned at about 5 to 7 weeks
of age [4].

Dispersal of juveniles - Juvenile coyotes usually disperse alone or
sometimes in groups at 6 to 9 months of age during October to February.
However, some juveniles do not disperse until their second year.
Juvenile coyotes may disperse up to 100 miles (160 km) from their den
[4]. In Minnesota, Berg and Chesness [7] reported mean dispersal
distances of 30 miles (48 km) that occurred at a mean rate of 7 miles
(11 km) per week [12]. Juvenile dispersal distances averaged 17 to 19
miles (28-31 km) in Alberta [48], 4 miles (7 km) in Arkansas [26], and 3
to 4 miles (5-6 km) in California [32].

Activity and movements - Coyotes are active day and night, with peaks in
activity at sunrise or sunset. Generally, activity and movements such
as foraging are greatest at night. Andelt [1] found that daytime
activity increased during the breeding season. In Arkansas, Gipson and
Sealander [26] found that young were more active than adults during the
day.

Life span - Coyotes in captivity may live as long as 18 years, but in
wild populations few coyotes live more than 6 to 8 years. The maximum
known age for a wild coyote is 14.5 years [4].
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Use of Fire in Population Management

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More info for the term: fire regime

Prescribed burning that favors small mammals by creating ecotones and
different age classes of vegetation would increase the prey base for
coyotes and make hunting easier by opening up the habitat [51].

FIRE REGIMES :
Find fire regime information for the plant communities in which this
species may occur by entering the species name in the FEIS home page under
"Find FIRE REGIMES".
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Tesky, Julie L. 1995. Canis latrans. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/

Coyote

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The coyote (Canis latrans) is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological niche as the golden jackal does in Eurasia. The coyote is larger and more predatory and was once referred to as the American jackal by a behavioral ecologist. Other historical names for the species include the prairie wolf and the brush wolf.

The coyote is listed as least concern by the International Union for Conservation of Nature, due to its wide distribution and abundance throughout North America. Coyote populations are also abundant southwards through Mexico and into Central America. The species is versatile, able to adapt to and expand into environments modified by humans. It is enlarging its range by moving into urban areas in the eastern U.S. and Canada. The coyote was sighted in eastern Panama (across the Panama Canal from their home range) for the first time in 2013.

The coyote has 19 recognized subspecies. The average male weighs 8 to 20 kg (18 to 44 lb) and the average female 7 to 18 kg (15 to 40 lb). Their fur color is predominantly light gray and red or fulvous interspersed with black and white, though it varies somewhat with geography. It is highly flexible in social organization, living either in a family unit or in loosely knit packs of unrelated individuals. Primarily carnivorous, its diet consists mainly of deer, rabbits, hares, rodents, birds, reptiles, amphibians, fish, and invertebrates, though it may also eat fruits and vegetables on occasion. Its characteristic vocalization is a howl made by solitary individuals. Humans are the coyote's greatest threat, followed by cougars and gray wolves. In spite of this, coyotes sometimes mate with gray, eastern, or red wolves, producing "coywolf" hybrids. In the northeastern regions of North America, the eastern coyote (a larger subspecies, though still smaller than wolves) is the result of various historical and recent matings with various types of wolves. Genetic studies show that most North American wolves contain some level of coyote DNA.

The coyote is a prominent character in Native American folklore, mainly in Aridoamerica, usually depicted as a trickster that alternately assumes the form of an actual coyote or a man. As with other trickster figures, the coyote uses deception and humor to rebel against social conventions. The animal was especially respected in Mesoamerican cosmology as a symbol of military might. After the European colonization of the Americas, it was seen in Anglo-American culture as a cowardly and untrustworthy animal. Unlike wolves, which have undergone an improvement of their public image, attitudes towards the coyote remain largely negative.

Description

 src=
Closeup of a mountain coyote's (C. l. lestes) head

Coyote males average 8 to 20 kg (18 to 44 lb) in weight, while females average 7 to 18 kg (15 to 40 lb), though size varies geographically. Northern subspecies, which average 18 kg (40 lb), tend to grow larger than the southern subspecies of Mexico, which average 11.5 kg (25 lb). Body length ranges on average from 1.0 to 1.35 m (3 ft 3 in to 4 ft 5 in), and tail length 40 cm (16 in), with females being shorter in both body length and height.[5] The largest coyote on record was a male killed near Afton, Wyoming, on November 19, 1937, which measured 1.5 m (4 ft 11 in) from nose to tail, and weighed 34 kg (75 lb).[6] Scent glands are located at the upper side of the base of the tail and are a bluish-black color.[7]

The color and texture of the coyote's fur vary somewhat geographically.[5] The hair's predominant color is light gray and red or fulvous, interspersed around the body with black and white. Coyotes living at high elevations tend to have more black and gray shades than their desert-dwelling counterparts, which are more fulvous or whitish-gray.[8] The coyote's fur consists of short, soft underfur and long, coarse guard hairs. The fur of northern subspecies is longer and denser than in southern forms, with the fur of some Mexican and Central American forms being almost hispid (bristly).[9] Generally, adult coyotes (including coywolf hybrids) have a sable coat color, dark neonatal coat color, bushy tail with an active supracaudal gland, and a white facial mask.[10] Albinism is extremely rare in coyotes; out of a total of 750,000 coyotes killed by federal and cooperative hunters between March 22, 1938, and June 30, 1945, only two were albinos.[8]

The coyote is typically smaller than the gray wolf, but has longer ears and a relatively larger braincase,[5] as well as a thinner frame, face, and muzzle. The scent glands are smaller than the gray wolf's, but are the same color.[7] Its fur color variation is much less varied than that of a wolf.[11] The coyote also carries its tail downwards when running or walking, rather than horizontally as the wolf does.[12]

Coyote tracks can be distinguished from those of dogs by their more elongated, less rounded shape.[13][14] Unlike dogs, the upper canines of coyotes extend past the mental foramina.[5]

Taxonomy and evolution

History

 src=
Toltec pictograph of a coyote

At the time of the European colonization of the Americas, coyotes were largely confined to open plains and arid regions of the western half of the continent.[15] In early post-Columbian historical records, determining whether the writer is describing coyotes or wolves is often difficult. One record from 1750 in Kaskaskia, Illinois, written by a local priest, noted that the "wolves" encountered there were smaller and less daring than European wolves. Another account from the early 1800s in Edwards County mentioned wolves howling at night, though these were likely coyotes.[16] This species was encountered several times during the Lewis and Clark Expedition (1804–1806), though it was already well known to European traders on the upper Missouri. Meriwether Lewis, writing on 5 May 1805, in northeastern Montana, described the coyote in these terms:

The small wolf or burrowing dog of the prairies are the inhabitants almost invariably of the open plains; they usually associate in bands of ten or twelve sometimes more and burrow near some pass or place much frequented by game; not being able alone to take deer or goat they are rarely ever found alone but hunt in bands; they frequently watch and seize their prey near their burrows; in these burrows, they raise their young and to them they also resort when pursued; when a person approaches them they frequently bark, their note being precisely that of the small dog. They are of an intermediate size between that of the fox and dog, very active fleet and delicately formed; the ears large erect and pointed the head long and pointed more like that of the fox; tale long ... the hair and fur also resembles the fox, tho' is much coarser and inferior. They are of a pale reddish-brown colour. The eye of a deep sea green colour small and piercing. Their [claws] are rather longer than those of the ordinary wolf or that common to the Atlantic states, none of which are to be found in this quarter, nor I believe above the river Plat.[17]

The coyote was first scientifically described by naturalist Thomas Say in September 1819, on the site of Lewis and Clark's Council Bluffs, 24 km (15 mi) up the Missouri River from the mouth of the Platte during a government-sponsored expedition with Major Stephen Long. He had the first edition of the Lewis and Clark journals in hand, which contained Biddle's edited version of Lewis's observations dated 5 May 1805. His account was published in 1823. Say was the first person to document the difference between a "prairie wolf" (coyote) and on the next page of his journal a wolf which he named Canis nubilus (Great Plains wolf).[3][18] Say described the coyote as:

Canis latrans. Cinereous or gray, varied with black above, and dull fulvous, or cinnamon; hair at base dusky plumbeous, in the middle of its length dull cinnamon, and at tip gray or black, longer on the vertebral line; ears erect, rounded at tip, cinnamon behind, the hair dark plumbeous at base, inside lined with gray hair; eyelids edged with black, superior eyelashes black beneath, and at tip above; supplemental lid margined with black-brown before, and edged with black brown behind; iris yellow; pupil black-blue; spot upon the lachrymal sac black-brown; rostrum cinnamon, tinctured with grayish on the nose; lips white, edged with black, three series of black seta; head between the ears intermixed with gray, and dull cinnamon, hairs dusky plumbeous at base; sides paler than the back, obsoletely fasciate with black above the legs; legs cinnamon on the outer side, more distinct on the posterior hair: a dilated black abbreviated line on the anterior ones near the wrist; tail bushy, fusiform, straight, varied with gray and cinnamon, a spot near the base above, and tip black; the tip of the trunk of the tail, attains the tip of the os calcis, when the leg is extended; beneath white, immaculate, tail cinnamon towards the tip, tip black; posterior feet four toed, anterior five toed.[3]

Naming and etymology

The earliest written reference to the species comes from the naturalist Francisco Hernández's Plantas y Animales de la Nueva España (1651), where it is described as a "Spanish fox" or "jackal". The first published usage of the word "coyote" (which is a Spanish borrowing of its Nahuatl name coyōtl About this soundpronunciation ) comes from the historian Francisco Javier Clavijero's Historia de México in 1780.[19] The first time it was used in English occurred in William Bullock's Six months' residence and travels in Mexico (1824), where it is variously transcribed as cayjotte and cocyotie. The word's spelling was standardized as "coyote" by the 1880s.[17][20] Alternative English names for the coyote include "prairie wolf", "brush wolf", "cased wolf",[21][a] "little wolf"[22] and "American jackal".[23] Its binomial name Canis latrans translates to "barking dog", a reference to the many vocalizations they produce.[24]

Evolution

Phylogenetic tree of the wolf-like canids with timing in millions of years[b] Caninae 3.5 Ma 3.0 2.5 2.0 0.96 0.6 0.38      

Domestic dog Tibetan mastiff (white background).jpg

   

Gray wolf Dogs, jackals, wolves, and foxes (Plate I).jpg

       

Coyote Dogs, jackals, wolves, and foxes (Plate IX).jpg

     

African golden wolf Dogs, jackals, wolves, and foxes (Plate XI).jpg

     

Golden jackal Dogs, jackals, wolves, and foxes (Plate X).jpg

     

Ethiopian wolf Dogs, jackals, wolves, and foxes (Plate VI).jpg

     

Dhole Dogs, jackals, wolves, and foxes (Plate XLI).jpg

     

African wild dog Dogs, jackals, wolves, and foxes (Plate XLIV).jpg

      2.6  

Side-striped jackal Dogs, jackals, wolves, and foxes (Plate XIII).jpg

   

Black-backed jackal Dogs, jackals, wolves, and foxes (Plate XII).jpg

       

Fossil record

 src=
Skeleton of Pleistocene coyote (C. l. orcutti)

Xiaoming Wang and Richard H. Tedford, one of the foremost authorities on carnivore evolution,[40] proposed that the genus Canis was the descendant of the coyote-like Eucyon davisi and its remains first appeared in the Miocene 6 million years ago (Mya) in the southwestern US and Mexico. By the Pliocene (5 Mya), the larger Canis lepophagus[41] appeared in the same region and by the early Pleistocene (1 Mya) C. latrans (the coyote) was in existence. They proposed that the progression from Eucyon davisi to C. lepophagus to the coyote was linear evolution.[42] Additionally, C. latrans and C. aureus are closely related to C. edwardii, a species that appeared earliest spanning the mid-Blancan (late Pliocene) to the close of the Irvingtonian (late Pleistocene), and coyote remains indistinguishable from C. latrans were contemporaneous with C. edwardii in North America.[43] Johnston describes C. lepophagus as having a more slender skull and skeleton than the modern coyote.[44] Ronald Nowak found that the early populations had small, delicate, narrowly proportioned skulls that resemble small coyotes and appear to be ancestral to C. latrans.[45]

C. lepophagus was similar in weight to modern coyotes, but had shorter limb bones that indicate a less cursorial lifestyle. The coyote represents a more primitive form of Canis than the gray wolf, as shown by its relatively small size and its comparatively narrow skull and jaws, which lack the grasping power necessary to hold the large prey in which wolves specialize. This is further corroborated by the coyote's sagittal crest, which is low or totally flattened, thus indicating a weaker bite than the wolves. The coyote is not a specialized carnivore as the wolf is, as shown by the larger chewing surfaces on the molars, reflecting the species' relative dependence on vegetable matter. In these respects, the coyote resembles the fox-like progenitors of the genus more so than the wolf.[46]

The oldest fossils that fall within the range of the modern coyote date to 0.74–0.85 Ma (million years) in Hamilton Cave, West Virginia; 0.73 Ma in Irvington, California; 0.35–0.48 Ma in Porcupine Cave, Colorado, and in Cumberland Cave, Pennsylvania.[47] Modern coyotes arose 1,000 years after the Quaternary extinction event.[48] Compared to their modern Holocene counterparts, Pleistocene coyotes (C. l. orcutti) were larger and more robust, likely in response to larger competitors and prey.[48] Pleistocene coyotes were likely more specialized carnivores than their descendants, as their teeth were more adapted to shearing meat, showing fewer grinding surfaces suited for processing vegetation.[49] Their reduction in size occurred within 1,000 years of the Quaternary extinction event, when their large prey died out.[48] Furthermore, Pleistocene coyotes were unable to exploit the big-game hunting niche left vacant after the extinction of the dire wolf (Aenocyon dirus), as it was rapidly filled by gray wolves, which likely actively killed off the large coyotes, with natural selection favoring the modern gracile morph.[49]

DNA evidence

In 1993, a study proposed that the wolves of North America display skull traits more similar to the coyote than wolves from Eurasia.[50] In 2010, a study found that the coyote was a basal member of the clade that included the Tibetan wolf, the domestic dog, the Mongolian wolf and the Eurasian wolf, with the Tibetan wolf diverging early from wolves and domestic dogs.[51] In 2016, a whole-genome DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago. The study also indicated that all North American wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry and that the red wolf and eastern wolf are highly admixed with different proportions of gray wolf and coyote ancestry.[52][53] The proposed timing of the wolf/coyote divergence conflicts with the finding of a coyote-like specimen in strata dated to 1 Mya.page needed]]]_56-0">page needed]]]-56">[54]

Genetic studies relating to wolves or dogs have inferred phylogenetic relationships based on the only reference genome available, that of the Boxer dog. In 2017, the first reference genome of the wolf Canis lupus lupus was mapped to aid future research.[55] In 2018, a study looked at the genomic structure and admixture of North American wolves, wolf-like canids, and coyotes using specimens from across their entire range that mapped the largest dataset of nuclear genome sequences against the wolf reference genome. The study supports the findings of previous studies that North American gray wolves and wolf-like canids were the result of complex gray wolf and coyote mixing. A polar wolf from Greenland and a coyote from Mexico represented the purest specimens. The coyotes from Alaska, California, Alabama, and Quebec show almost no wolf ancestry. Coyotes from Missouri, Illinois, and Florida exhibit 5–10% wolf ancestry. There was 40%:60% wolf to coyote ancestry in red wolves, 60%:40% in Eastern timber wolves, and 75%:25% in the Great Lakes wolves. There was 10% coyote ancestry in Mexican wolves and the Atlantic Coast wolves, 5% in Pacific Coast and Yellowstone wolves, and less than 3% in Canadian archipelago wolves. If a third canid had been involved in the admixture of the North American wolf-like canids then its genetic signature would have been found in coyotes and wolves, which it has not.[56]

In 2018, whole genome sequencing was used to compare members of the genus Canis. The study indicates that the common ancestor of the coyote and gray wolf has genetically admixed with a ghost population of an extinct unidentified canid. The canid was genetically close to the dhole and had evolved after the divergence of the African wild dog from the other canid species. The basal position of the coyote compared to the wolf is proposed to be due to the coyote retaining more of the mitochondrial genome of this unknown canid.[57]

Subspecies

As of 2005, 19 subspecies are recognized.[23][58] Geographic variation in coyotes is not great, though taken as a whole, the eastern subspecies (C. l. thamnos and C. l. frustor) are large, dark-colored animals, with a gradual paling in color and reduction in size westward and northward (C. l. texensis, C. l. latrans, C. l. lestes, and C. l. incolatus), a brightening of ochraceous tones – deep orange or brown – towards the Pacific coast (C. l. ochropus, C. l. umpquensis), a reduction in size in Aridoamerica (C. l. microdon, C. l. mearnsi) and a general trend towards dark reddish colors and short muzzles in Mexican and Central American populations.[59]

Hybridization

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Melanistic coyotes owe their color to a mutation that first arose in domestic dogs.[71]

Coyotes have occasionally mated with domestic dogs, sometimes producing crosses colloquially known as "coydogs".[72] Such matings are rare in the wild, as the mating cycles of dogs and coyotes do not coincide, and coyotes are usually antagonistic towards dogs. Hybridization usually only occurs when coyotes are expanding into areas where conspecifics are few, and dogs are the only alternatives. Even then, pup survival rates are lower than normal, as dogs do not form pair bonds with coyotes, thus making the rearing of pups more difficult.[73] In captivity, F1 hybrids (first generation) tend to be more mischievous and less manageable as pups than dogs, and are less trustworthy on maturity than wolf-dog hybrids.[72] Hybrids vary in appearance, but generally retain the coyote's usual characteristics. F1 hybrids tend to be intermediate in form between dogs and coyotes, while F2 hybrids (second generation) are more varied. Both F1 and F2 hybrids resemble their coyote parents in terms of shyness and intrasexual aggression.[10][74] Hybrids are fertile and can be successfully bred through four generations.[72] Melanistic coyotes owe their black pelts to a mutation that first arose in domestic dogs.[71] A population of nonalbino white coyotes in Newfoundland owe their coloration to a melanocortin 1 receptor mutation inherited from Golden Retrievers.[75]

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A coywolf hybrid conceived in captivity between a male gray wolf and a female coyote

Coyotes have hybridized with wolves to varying degrees, particularly in eastern North America. The so-called "eastern coyote" of northeastern North America probably originated in the aftermath of the extermination of gray and eastern wolves in the northeast, thus allowing coyotes to colonize former wolf ranges and mix with the remnant wolf populations. This hybrid is smaller than either the gray or eastern wolf, and holds smaller territories, but is in turn larger and holds more extensive home ranges than the typical western coyote. As of 2010, the eastern coyote's genetic makeup is fairly uniform, with minimal influence from eastern wolves or western coyotes.[76] Adult eastern coyotes are larger than western coyotes, with female eastern coyotes weighing 21% more than male western coyotes.[76][77] Physical differences become more apparent by the age of 35 days, with eastern coyote pups having longer legs than their western counterparts. Differences in dental development also occurs, with tooth eruption being later, and in a different order in the eastern coyote.[78] Aside from its size, the eastern coyote is physically similar to the western coyote. The four color phases range from dark brown to blond or reddish blond, though the most common phase is gray-brown, with reddish legs, ears, and flanks.[79] No significant differences exist between eastern and western coyotes in aggression and fighting, though eastern coyotes tend to fight less, and are more playful. Unlike western coyote pups, in which fighting precedes play behavior, fighting among eastern coyote pups occurs after the onset of play.[78] Eastern coyotes tend to reach sexual maturity at two years of age, much later than in western coyotes.[76]

Eastern and red wolves are also products of varying degrees of wolf-coyote hybridization. The eastern wolf probably was a result of a wolf-coyote admixture, combined with extensive backcrossing with parent gray wolf populations. The red wolf may have originated during a time of declining wolf populations in the Southeastern Woodlands, forcing a wolf-coyote hybridization, as well as backcrossing with local parent coyote populations to the extent that about 75–80% of the modern red wolf's genome is of coyote derivation.[52][80]

Behavior

Social and reproductive behaviors

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Mearns' coyote (C. l. mearnsi) pups playing
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A pack of coyotes in Yellowstone National Park

Like the Eurasian golden jackal, the coyote is gregarious, but not as dependent on conspecifics as more social canid species like wolves are. This is likely because the coyote is not a specialized hunter of large prey as the latter species is.[81] The basic social unit of a coyote pack is a family containing a reproductive female. However, unrelated coyotes may join forces for companionship, or to bring down prey too large to attack singly. Such "nonfamily" packs are only temporary, and may consist of bachelor males, nonreproductive females and subadult young. Families are formed in midwinter, when females enter estrus.[22] Pair bonding can occur 2–3 months before actual copulation takes place.[82] The copulatory tie can last 5–45 minutes.[83] A female entering estrus attracts males by scent marking[84] and howling with increasing frequency.[23] A single female in heat can attract up to seven reproductive males, which can follow her for as long as a month. Although some squabbling may occur among the males, once the female has selected a mate and copulates, the rejected males do not intervene, and move on once they detect other estrous females.[22] Unlike the wolf, which has been known to practice both monogamous and bigamous matings,[85] the coyote is strictly monogamous, even in areas with high coyote densities and abundant food.[86] Females that fail to mate sometimes assist their sisters or mothers in raising their pups, or join their siblings until the next time they can mate. The newly mated pair then establishes a territory and either constructs their own den or cleans out abandoned badger, marmot, or skunk earths. During the pregnancy, the male frequently hunts alone and brings back food for the female. The female may line the den with dried grass or with fur pulled from her belly.[22] The gestation period is 63 days, with an average litter size of six, though the number fluctuates depending on coyote population density and the abundance of food.[23]

Coyote pups are born in dens, hollow trees, or under ledges, and weigh 200 to 500 g (0.44 to 1.10 lb) at birth. They are altricial, and are completely dependent on milk for their first 10 days. The incisors erupt at about 12 days, the canines at 16, and the second premolars at 21. Their eyes open after 10 days, by which point the pups become increasingly more mobile, walking by 20 days, and running at the age of six weeks. The parents begin supplementing the pup's diet with regurgitated solid food after 12–15 days. By the age of four to six weeks, when their milk teeth are fully functional, the pups are given small food items such as mice, rabbits, or pieces of ungulate carcasses, with lactation steadily decreasing after two months.[22] Unlike wolf pups, coyote pups begin seriously fighting (as opposed to play fighting) prior to engaging in play behavior. A common play behavior includes the coyote "hip-slam".[74] By three weeks of age, coyote pups bite each other with less inhibition than wolf pups. By the age of four to five weeks, pups have established dominance hierarchies, and are by then more likely to play rather than fight.[87] The male plays an active role in feeding, grooming, and guarding the pups, but abandons them if the female goes missing before the pups are completely weaned. The den is abandoned by June to July, and the pups follow their parents in patrolling their territory and hunting. Pups may leave their families in August, though can remain for much longer. The pups attain adult dimensions at eight months and gain adult weight a month later.[22]

Territorial and sheltering behaviors

Individual feeding territories vary in size from 0.4 to 62 km2 (0.15 to 24 sq mi), with the general concentration of coyotes in a given area depending on food abundance, adequate denning sites, and competition with conspecifics and other predators. The coyote generally does not defend its territory outside of the denning season,[22] and is much less aggressive towards intruders than the wolf is, typically chasing and sparring with them, but rarely killing them.[88] Conflicts between coyotes can arise during times of food shortage.[22] Coyotes mark their territories by raised-leg urination and ground-scratching.[89][84]

Like wolves, coyotes use a den (usually the deserted holes of other species) when gestating and rearing young, though they may occasionally give birth under sagebrushes in the open. Coyote dens can be located in canyons, washouts, coulees, banks, rock bluffs, or level ground. Some dens have been found under abandoned homestead shacks, grain bins, drainage pipes, railroad tracks, hollow logs, thickets, and thistles. The den is continuously dug and cleaned out by the female until the pups are born. Should the den be disturbed or infested with fleas, the pups are moved into another den. A coyote den can have several entrances and passages branching out from the main chamber.[90] A single den can be used year after year.[23]

Hunting and feeding behaviors

While the popular consensus is that olfaction is very important for hunting,[91] two studies that experimentally investigated the role of olfactory, auditory, and visual cues found that visual cues are the most important ones for hunting in red foxes[92] and coyotes.[93][94]

A coyote is pouncing.
A coyote pouncing on prey.

When hunting large prey, the coyote often works in pairs or small groups.[5] Success in killing large ungulates depends on factors such as snow depth and crust density. Younger animals usually avoid participating in such hunts, with the breeding pair typically doing most of the work.[23] Unlike the wolf, which attacks large prey from the rear, the coyote approaches from the front, lacerating its prey's head and throat. Like other canids, the coyote caches excess food.[95] Coyotes catch mouse-sized rodents by pouncing, whereas ground squirrels are chased. Although coyotes can live in large groups, small prey is typically caught singly.[23] Coyotes have been observed to kill porcupines in pairs, using their paws to flip the rodents on their backs, then attacking the soft underbelly. Only old and experienced coyotes can successfully prey on porcupines, with many predation attempts by young coyotes resulting in them being injured by their prey's quills.[96] Coyotes sometimes urinate on their food, possibly to claim ownership over it.[89][97] Recent evidence demonstrates that at least some coyotes have become more nocturnal in hunting, presumably to avoid humans.[98]

Coyotes may occasionally form mutualistic hunting relationships with American badgers, assisting each other in digging up rodent prey.[99] The relationship between the two species may occasionally border on apparent "friendship", as some coyotes have been observed laying their heads on their badger companions or licking their faces without protest. The amicable interactions between coyotes and badgers were known to pre-Columbian civilizations, as shown on a Mexican jar dated to 1250–1300 CE depicting the relationship between the two.[100]

Food scraps, pet food, and animal feces may attract a coyote to a trash can.[101]

Communication

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A coyote howling

Body language

Being both a gregarious and solitary animal, the variability of the coyote's visual and vocal repertoire is intermediate between that of the solitary foxes and the highly social wolf.[81] The aggressive behavior of the coyote bears more similarities to that of foxes than it does that of wolves and dogs. An aggressive coyote arches its back and lowers its tail.[102] Unlike dogs, which solicit playful behavior by performing a "play-bow" followed by a "play-leap", play in coyotes consists of a bow, followed by side-to-side head flexions and a series of "spins" and "dives". Although coyotes will sometimes bite their playmates' scruff as dogs do, they typically approach low, and make upward-directed bites.[103] Pups fight each other regardless of sex, while among adults, aggression is typically reserved for members of the same sex. Combatants approach each other waving their tails and snarling with their jaws open, though fights are typically silent. Males tend to fight in a vertical stance, while females fight on all four paws. Fights among females tend to be more serious than ones among males, as females seize their opponents' forelegs, throat, and shoulders.[102]

Vocalizations

A yelping coyote

The coyote has been described as "the most vocal of all [wild] North American mammals".[104][105] Its loudness and range of vocalizations was the cause for its binomial name Canis latrans, meaning "barking dog". At least 11 different vocalizations are known in adult coyotes. These sounds are divided into three categories: agonistic and alarm, greeting, and contact. Vocalizations of the first category include woofs, growls, huffs, barks, bark howls, yelps, and high-frequency whines. Woofs are used as low-intensity threats or alarms and are usually heard near den sites, prompting the pups to immediately retreat into their burrows. Growls are used as threats at short distances but have also been heard among pups playing and copulating males. Huffs are high-intensity threat vocalizations produced by rapid expiration of air. Barks can be classed as both long-distance threat vocalizations and alarm calls. Bark howls may serve similar functions. Yelps are emitted as a sign of submission, while high-frequency whines are produced by dominant animals acknowledging the submission of subordinates. Greeting vocalizations include low-frequency whines, 'wow-oo-wows', and group yip howls. Low-frequency whines are emitted by submissive animals and are usually accompanied by tail wagging and muzzle nibbling. The sound known as 'wow-oo-wow' has been described as a "greeting song". The group yip howl is emitted when two or more pack members reunite and may be the final act of a complex greeting ceremony. Contact calls include lone howls and group howls, as well as the previously mentioned group yip howls. The lone howl is the most iconic sound of the coyote and may serve the purpose of announcing the presence of a lone individual separated from its pack. Group howls are used as both substitute group yip howls and as responses to either lone howls, group howls, or group yip howls.[24]

Ecology

Habitat

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An urban coyote in Bernal Heights, San Francisco

Prior to the near extermination of wolves and cougars, the coyote was most numerous in grasslands inhabited by bison, pronghorn, elk, and other deer, doing particularly well in short-grass areas with prairie dogs, though it was just as much at home in semiarid areas with sagebrush and jackrabbits or in deserts inhabited by cactus, kangaroo rats, and rattlesnakes. As long as it was not in direct competition with the wolf, the coyote ranged from the Sonoran Desert to the alpine regions of adjoining mountains or the plains and mountainous areas of Alberta. With the extermination of the wolf, the coyote's range expanded to encompass broken forests from the tropics of Guatemala and the northern slope of Alaska.[22]

Coyotes walk around 5–16 kilometres (3–10 mi) per day, often along trails such as logging roads and paths; they may use iced-over rivers as travel routes in winter. They are often crepuscular, being more active around evening and the beginning of the night than during the day. Like many canids, coyotes are competent swimmers, reported to be able to travel at least 0.8 kilometres (0.5 mi) across water.[106]

Diet

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A coyote with a scrap of road-killed pronghorn in Seedskadee National Wildlife Refuge, Wyoming

The coyote is ecologically the North American equivalent of the Eurasian golden jackal.[107] Likewise, the coyote is highly versatile in its choice of food, but is primarily carnivorous, with 90% of its diet consisting of meat. Prey species include bison (largely as carrion), white-tailed deer, mule deer, moose, elk, bighorn sheep, pronghorn, rabbits, hares, rodents, birds (especially galliformes, young water birds and pigeons and doves), amphibians (except toads), lizards, snakes, turtles and tortoises, fish, crustaceans, and insects. Coyotes may be picky over the prey they target, as animals such as shrews, moles, and brown rats do not occur in their diet in proportion to their numbers.[22] However, terrestrial and/or burrowing small mammals such as ground squirrels and associated species (marmots, prairie dogs, chipmunks) as well as voles, pocket gophers, kangaroo rats and other ground-favoring rodents may be quite common foods, especially for lone coyotes.[108][109][110] More unusual prey include fishers,[111] young black bear cubs,[112] harp seals[113] and rattlesnakes. Coyotes kill rattlesnakes mostly for food (but also to protect their pups at their dens) by teasing the snakes until they stretch out and then biting their heads and snapping and shaking the snakes.[114] Birds taken by coyotes may range in size from thrashers, larks and sparrows to adult wild turkeys and, possibly, brooding adult swans and pelicans.[115][116][117][118] If working in packs or pairs, coyotes may have access to larger prey than lone individuals normally take, such as various prey weighing more than 10 kg (22 lb).[119][120] In some cases, packs of coyotes have dispatched much larger prey such as adult Odocoileus deer, cow elk, pronghorns and wild sheep, although the young fawn, calves and lambs of these animals are considerably more often taken even by packs, as well as domestic sheep and domestic cattle. In some cases, coyotes can bring down prey weighing up to 100 to 200 kg (220 to 440 lb) or more. When it comes to adult ungulates such as wild deer, they often exploit them when vulnerable such as those that are infirm, stuck in snow or ice, otherwise winter-weakened or heavily pregnant, whereas less wary domestic ungulates may be more easily exploited.[119][121][122][123][124][125][126]

Although coyotes prefer fresh meat, they will scavenge when the opportunity presents itself. Excluding the insects, fruit, and grass eaten, the coyote requires an estimated 600 g (1.3 lb) of food daily, or 250 kg (550 lb) annually.[22] The coyote readily cannibalizes the carcasses of conspecifics, with coyote fat having been successfully used by coyote hunters as a lure or poisoned bait.[7] The coyote's winter diet consists mainly of large ungulate carcasses, with very little plant matter. Rodent prey increases in importance during the spring, summer, and fall.[5]

The coyote feeds on a variety of different produce, including blackberries, blueberries, peaches, pears, apples, prickly pears, chapotes, persimmons, peanuts, watermelons, cantaloupes, and carrots. During the winter and early spring, the coyote eats large quantities of grass, such as green wheat blades. It sometimes eats unusual items such as cotton cake, soybean meal, domestic animal droppings, beans, and cultivated grain such as maize, wheat, and sorghum.[22]

In coastal California, coyotes now consume a higher percentage of marine-based food than their ancestors, which is thought to be due to the extirpation of the grizzly bear from this region.[127] In Death Valley, coyotes may consume great quantities of hawkmoth caterpillars or beetles in the spring flowering months.[128]

Enemies and competitors

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Comparative illustration of coyote and gray wolf
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Mountain coyotes (C. l. lestes) cornering a juvenile cougar

In areas where the ranges of coyotes and gray wolves overlap, interference competition and predation by wolves has been hypothesized to limit local coyote densities. Coyote ranges expanded during the 19th and 20th centuries following the extirpation of wolves, while coyotes were driven to extinction on Isle Royale after wolves colonized the island in the 1940s. One study conducted in Yellowstone National Park, where both species coexist, concluded that the coyote population in the Lamar River Valley declined by 39% following the reintroduction of wolves in the 1990s, while coyote populations in wolf inhabited areas of the Grand Teton National Park are 33% lower than in areas where they are absent.[129][130] Wolves have been observed to not tolerate coyotes in their vicinity, though coyotes have been known to trail wolves to feed on their kills.[100]

Coyotes may compete with cougars in some areas. In the eastern Sierra Nevada, coyotes compete with cougars over mule deer. Cougars normally outcompete and dominate coyotes, and may kill them occasionally, thus reducing coyote predation pressure on smaller carnivores such as foxes and bobcats.[131] Coyotes that are killed are sometimes not eaten, perhaps indicating that these comprise competitive interspecies interactions, however there are multiple confirmed cases of cougars also eating coyotes.[132][133] In northeastern Mexico, cougar predation on coyotes continues apace but coyotes were absent from the prey spectrum of sympatric jaguars, apparently due to differing habitat usages.[134]

Other than by gray wolves and cougars, predation on adult coyotes is relatively rare but multiple other predators can be occasional threats. In some cases, adult coyotes have been preyed upon by both American black and grizzly bears,[135] American alligators,[136] large Canada lynx[137] and golden eagles.[138] At kill sites and carrion, coyotes, especially if working alone, tend to be dominated by wolves, cougars, bears, wolverines and, usually but not always, eagles (i.e., bald and golden). When such larger, more powerful and/or more aggressive predators such as these come to a shared feeding site, a coyote may either try to fight, wait until the other predator is done or occasionally share a kill, but if a major danger such as wolves or an adult cougar is present, the coyote will tend to flee.[139][140][141][142][143][144][145][146]

Coyotes rarely kill healthy adult red foxes, and have been observed to feed or den alongside them, though they often kill foxes caught in traps. Coyotes may kill fox kits, but this is not a major source of mortality.[147] In southern California, coyotes frequently kill gray foxes, and these smaller canids tend to avoid areas with high coyote densities.[148]

In some areas, coyotes share their ranges with bobcats. These two similarly-sized species rarely physically confront one another, though bobcat populations tend to diminish in areas with high coyote densities.[149] However, several studies have demonstrated interference competition between coyotes and bobcats, and in all cases coyotes dominated the interaction.[150][151] Multiple researchers[152][153][154][151][155] reported instances of coyotes killing bobcats, whereas bobcats killing coyotes is more rare.[150] Coyotes attack bobcats using a bite-and-shake method similar to what is used on medium-sized prey. Coyotes (both single individuals and groups) have been known to occasionally kill bobcats – in most cases, the bobcats were relatively small specimens, such as adult females and juveniles.[151] However, coyote attacks (by an unknown number of coyotes) on adult male bobcats have occurred. In California, coyote and bobcat populations are not negatively correlated across different habitat types, but predation by coyotes is an important source of mortality in bobcats.[148] Biologist Stanley Paul Young noted that in his entire trapping career, he had never successfully saved a captured bobcat from being killed by coyotes, and wrote of two incidents wherein coyotes chased bobcats up trees.[100] Coyotes have been documented to directly kill Canada lynx on occasion,[156][157][158] and compete with them for prey, especially snowshoe hares.[156] In some areas, including central Alberta, lynx are more abundant where coyotes are few, thus interactions with coyotes appears to influence lynx populations more than the availability of snowshoe hares.[159]

Range

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Range of coyote subspecies as of 1978: (1) Mexican coyote, (2) San Pedro Martir coyote, (3) El Salvador coyote, (4) southeastern coyote, (5) Belize coyote, (6) Honduras coyote, (7) Durango coyote, (8) northern coyote, (9) Tiburón Island coyote, (10) plains coyote, (11) mountain coyote, (12) Mearns' coyote, (13) Lower Rio Grande coyote, (14) California valley coyote, (15) peninsula coyote, (16) Texas plains coyote, (17) northeastern coyote, (18) northwest coast coyote, (19) Colima coyote, (20) eastern coyote[61]
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Coyote expansion over the past 10,000 years[160]
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Coyote expansion over the decades since 1900[160]

Due to the coyote's wide range and abundance throughout North America, it is listed as Least Concern by the International Union for Conservation of Nature (IUCN).[2] The coyote's pre-Columbian range was limited to the Southwest and Plains regions of North America, and northern and central Mexico. By the 19th century, the species expanded north and east, expanding further after 1900, coinciding with land conversion and the extirpation of wolves. By this time, its range encompassed the entire North American continent, including all of the contiguous United States and Mexico, southward into Central America, and northward into most of Canada and Alaska.[161] This expansion is ongoing, and the species now occupies the majority of areas between 8°N (Panama) and 70°N (northern Alaska).[2]

Although it was once widely believed that coyotes are recent immigrants to southern Mexico and Central America, aided in their expansion by deforestation, Pleistocene and Early Holocene records, as well as records from the pre-Columbian period and early European colonization show that the animal was present in the area long before modern times. Nevertheless, range expansion did occur south of Costa Rica during the late 1970s and northern Panama in the early 1980s, following the expansion of cattle-grazing lands into tropical rain forests. The coyote is predicted to appear in northern Belize in the near future, as the habitat there is favorable to the species.[162] Concerns have been raised of a possible expansion into South America through the Panamanian Isthmus, should the Darién Gap ever be closed by the Pan-American Highway.[163] This fear was partially confirmed in January 2013, when the species was recorded in eastern Panama's Chepo District, beyond the Panama Canal.[64]

A 2017 genetic study proposes that coyotes were originally not found in the area of the eastern United States. From the 1890s, dense forests were transformed into agricultural land and wolf control implemented on a large scale, leaving a niche for coyotes to disperse into. There were two major dispersals from two populations of genetically distinct coyotes. The first major dispersal to the northeast came in the early 20th century from those coyotes living in the northern Great Plains. These came to New England via the northern Great Lakes region and southern Canada, and to Pennsylvania via the southern Great Lakes region, meeting together in the 1940s in New York and Pennsylvania. These coyotes have hybridized with the remnant gray wolf and eastern wolf populations, which has added to coyote genetic diversity and may have assisted adaptation to the new niche. The second major dispersal to the southeast came in the mid-20th century from Texas and reached the Carolinas in the 1980s. These coyotes have hybridized with the remnant red wolf populations before the 1970s when the red wolf was extirpated in the wild, which has also added to coyote genetic diversity and may have assisted adaptation to this new niche as well. Both of these two major coyote dispersals have experienced rapid population growth and are forecast to meet along the mid-Atlantic coast. The study concludes that for coyotes the long range dispersal, gene flow from local populations, and rapid population growth may be inter-related.[164]

In July 2018 Cambridge, Ontario city government removed leg traps from a city park after complaints about perceived harm to the coyotes.[165][166]

Diseases and parasites

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California valley coyote (C. l. ochropus) suffering from sarcoptic mange

Among large North American carnivores, the coyote probably carries the largest number of diseases and parasites, likely due to its wide range and varied diet.[167] Viral diseases known to infect coyotes include rabies, canine distemper, infectious canine hepatitis, four strains of equine encephalitis, and oral papillomatosis. By the late 1970s, serious rabies outbreaks in coyotes had ceased to be a problem for over 60 years, though sporadic cases every 1–5 years did occur. Distemper causes the deaths of many pups in the wild, though some specimens can survive infection. Tularemia, a bacterial disease, infects coyotes from tick bites and through their rodent and lagomorph prey, and can be deadly for pups.[168]

Coyotes can be infected by both demodectic and sarcoptic mange, the latter being the most common. Mite infestations are rare and incidental in coyotes, while tick infestations are more common, with seasonal peaks depending on locality (May–August in the Northwest, March–November in Arkansas). Coyotes are only rarely infested with lice, while fleas infest coyotes from puphood, though they may be more a source of irritation than serious illness. Pulex simulans is the most common species to infest coyotes, while Ctenocephalides canis tends to occur only in places where coyotes and dogs (its primary host) inhabit the same area. Although coyotes are rarely host to flukes, they can nevertheless have serious effects on coyotes, particularly Nanophyetus salmincola, which can infect them with salmon poisoning disease, a disease with a 90% mortality rate. Trematode Metorchis conjunctus can also infect coyotes.[169] Tapeworms have been recorded to infest 60–95% of all coyotes examined. The most common species to infest coyotes are Taenia pisiformis and Taenia crassiceps, which uses cottontail rabbits as intermediate hosts. The largest species known in coyotes is T. hydatigena, which enters coyotes through infected ungulates, and can grow to lengths of 80 to 400 cm (31 to 157 in). Although once largely limited to wolves, Echinococcus granulosus has expanded to coyotes since the latter began colonizing former wolf ranges. The most frequent ascaroid roundworm in coyotes is Toxascaris leonina, which dwells in the coyote's small intestine and has no ill effects, except for causing the host to eat more frequently. Hookworms of the genus Ancylostoma infest coyotes throughout their range, being particularly prevalent in humid areas. In areas of high moisture, such as coastal Texas, coyotes can carry up to 250 hookworms each. The blood-drinking A. caninum is particularly dangerous, as it damages the coyote through blood loss and lung congestion. A 10-day-old pup can die from being host to as few as 25 A. caninum worms.[168]

Relationships with humans

In folklore and mythology

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Coyote paddling in a canoe in Edward S. Curtis's Indian days of long ago

Coyote features as a trickster figure and skin-walker in the folktales of some Native Americans, notably several nations in the Southwestern and Plains regions, where he alternately assumes the form of an actual coyote or that of a man. As with other trickster figures, Coyote acts as a picaresque hero who rebels against social convention through deception and humor.[170] Folklorists such as Harris believe coyotes came to be seen as tricksters due to the animal's intelligence and adaptability.[171] After the European colonization of the Americas, Anglo-American depictions of Coyote are of a cowardly and untrustworthy animal.[172] Unlike the gray wolf, which has undergone a radical improvement of its public image, Anglo-American cultural attitudes towards the coyote remain largely negative.[173]

In the Maidu creation story, Coyote introduces work, suffering, and death to the world. Zuni lore has Coyote bringing winter into the world by stealing light from the kachinas. The Chinook, Maidu, Pawnee, Tohono O'odham, and Ute portray the coyote as the companion of The Creator. A Tohono O'odham flood story has Coyote helping Montezuma survive a global deluge that destroys humanity. After The Creator creates humanity, Coyote and Montezuma teach people how to live. The Crow creation story portrays Old Man Coyote as The Creator. In The Dineh creation story, Coyote was present in the First World with First Man and First Woman, though a different version has it being created in the Fourth World. The Navajo Coyote brings death into the world, explaining that without death, too many people would exist, thus no room to plant corn.[174]

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Mural from Atetelco, Teotihuacán depicting coyote warriors

Prior to the Spanish conquest of the Aztec Empire, Coyote played a significant role in Mesoamerican cosmology. The coyote symbolized military might in Classic era Teotihuacan, with warriors dressing up in coyote costumes to call upon its predatory power. The species continued to be linked to Central Mexican warrior cults in the centuries leading up to the post-Classic Aztec rule.[175] In Aztec mythology, Huehuecóyotl (meaning "old coyote"), the god of dance, music and carnality, is depicted in several codices as a man with a coyote's head.[176] He is sometimes depicted as a womanizer, responsible for bringing war into the world by seducing Xochiquetzal, the goddess of love.[177] Epigrapher David H. Kelley argued that the god Quetzalcoatl owed its origins to pre-Aztec Uto-Aztecan mythological depictions of the coyote, which is portrayed as mankind's "Elder Brother", a creator, seducer, trickster, and culture hero linked to the morning star.[178]

Attacks on humans

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A sign discouraging people from feeding coyotes, which can lead to them habituating themselves to human presence, thus increasing the likelihood of attacks

Coyote attacks on humans are uncommon and rarely cause serious injuries, due to the relatively small size of the coyote, but have been increasingly frequent, especially in California. There have been only two confirmed fatal attacks: one on a three-year-old named Kelly Keen in Glendale, California[179] and another on a nineteen-year-old named Taylor Mitchell in Nova Scotia, Canada.[180] In the 30 years leading up to March 2006, at least 160 attacks occurred in the United States, mostly in the Los Angeles County area.[181] Data from United States Department of Agriculture (USDA) Wildlife Services, the California Department of Fish and Game, and other sources show that while 41 attacks occurred during the period of 1988–1997, 48 attacks were verified from 1998 through 2003. The majority of these incidents occurred in Southern California near the suburban-wildland interface.[179]

In the absence of the harassment of coyotes practiced by rural people, urban coyotes are losing their fear of humans, which is further worsened by people intentionally or unintentionally feeding coyotes. In such situations, some coyotes have begun to act aggressively toward humans, chasing joggers and bicyclists, confronting people walking their dogs, and stalking small children.[179] Non-rabid coyotes in these areas sometimes target small children, mostly under the age of 10, though some adults have been bitten.[182]

Although media reports of such attacks generally identify the animals in question as simply "coyotes", research into the genetics of the eastern coyote indicates those involved in attacks in northeast North America, including Pennsylvania, New York, New England, and eastern Canada, may have actually been coywolves, hybrids of Canis latrans and C. lupus, not fully coyotes.[183]

Livestock and pet predation

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Coyote confronting a dog

As of 2007, coyotes were the most abundant livestock predators in western North America, causing the majority of sheep, goat, and cattle losses.[184] For example, according to the National Agricultural Statistics Service, coyotes were responsible for 60.5% of the 224,000 sheep deaths attributed to predation in 2004.[185][186] The total number of sheep deaths in 2004 comprised 2.22% of the total sheep and lamb population in the United States,[187] which, according to the National Agricultural Statistics Service USDA report, totaled 4.66 million and 7.80 million heads respectively as of July 1, 2005.[188] Because coyote populations are typically many times greater and more widely distributed than those of wolves, coyotes cause more overall predation losses. United States government agents routinely shoot, poison, trap, and kill about 90,000 coyotes each year to protect livestock.[189] An Idaho census taken in 2005 showed that individual coyotes were 5% as likely to attack livestock as individual wolves.[190] In Utah, more than 11,000 coyotes were killed for bounties totaling over $500,000 in the fiscal year ending June 30, 2017.[191]

Livestock guardian dogs are commonly used to aggressively repel predators and have worked well in both fenced pasture and range operations.[192] A 1986 survey of sheep producers in the USA found that 82% reported the use of dogs represented an economic asset.[193]

Re-wilding cattle, which involves increasing the natural protective tendencies of cattle, is a method for controlling coyotes discussed by Temple Grandin of Colorado State University.[194] This method is gaining popularity among producers who allow their herds to calve on the range and whose cattle graze open pastures throughout the year.[195]

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Coyote with a typical throat hold on a domestic sheep

Coyotes typically bite the throat just behind the jaw and below the ear when attacking adult sheep or goats, with death commonly resulting from suffocation. Blood loss is usually a secondary cause of death. Calves and heavily fleeced sheep are killed by attacking the flanks or hindquarters, causing shock and blood loss. When attacking smaller prey, such as young lambs, the kill is made by biting the skull and spinal regions, causing massive tissue and bone damage. Small or young prey may be completely carried off, leaving only blood as evidence of a kill. Coyotes usually leave the hide and most of the skeleton of larger animals relatively intact, unless food is scarce, in which case they may leave only the largest bones. Scattered bits of wool, skin, and other parts are characteristic where coyotes feed extensively on larger carcasses.[184]

Tracks are an important factor in distinguishing coyote from dog predation. Coyote tracks tend to be more oval-shaped and compact than those of domestic dogs, and their claw marks are less prominent and the tracks tend to follow a straight line more closely than those of dogs. With the exception of sighthounds, most dogs of similar weight to coyotes have a slightly shorter stride.[184] Coyote kills can be distinguished from wolf kills by less damage to the underlying tissues in the former. Also, coyote scat tends to be smaller than wolf scat.[196][197]

Coyotes are often attracted to dog food and animals that are small enough to appear as prey. Items such as garbage, pet food, and sometimes feeding stations for birds and squirrels attract coyotes into backyards. About three to five pets attacked by coyotes are brought into the Animal Urgent Care hospital of South Orange County (California) each week, the majority of which are dogs, since cats typically do not survive the attacks.[198] Scat analysis collected near Claremont, California, revealed that coyotes relied heavily on pets as a food source in winter and spring.[179] At one location in Southern California, coyotes began relying on a colony of feral cats as a food source. Over time, the coyotes killed most of the cats and then continued to eat the cat food placed daily at the colony site by people who were maintaining the cat colony.[179] Coyotes usually attack smaller-sized dogs, but they have been known to attack even large, powerful breeds such as the Rottweiler in exceptional cases.[199] Dogs larger than coyotes, such as greyhounds, are generally able to drive them off and have been known to kill coyotes.[200] Smaller breeds are more likely to suffer injury or death.[182]

Hunting

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Coyote tracks compared to those of the domestic dog

Coyote hunting is one of the most common forms of predator hunting that humans partake in. There are not many regulations with regard to the taking of the coyote which means there are many different methods that can be used to hunt the animal. The most common forms are trapping, calling, and hound hunting.[201] Since coyotes are colorblind, seeing only in shades of gray and subtle blues, open camouflages, and plain patterns can be used. The average male coyote weighs 8 to 20 kg (18 to 44 lbs) and the average female coyote 7 to 18 kg (15 to 40 lbs) a universal projectile that can perform between those weights is the .223 Remington so that the projectile expands in the target after the entry but before the exit thus delivering the most energy.[202] Coyotes being the light and agile animals they are, they often leave a very light impression on terrain. The coyote's footprint is oblong, approximately 6.35 cm (2.5-inches) long and 5.08 cm (2-inches) wide. There are 4 claws in both their front and hind paws. The coyote's center pad is relatively shaped like that of a rounded triangle. Like the domestic dog the coyote's front paw is slightly larger than the hind paw. The coyote's paw is most similar to that of the domestic dog.[203]

Uses

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Fur of a Canadian coyote

Prior to the mid-19th century, coyote fur was considered worthless. This changed with the diminution of beavers, and by 1860, the hunting of coyotes for their fur became a great source of income (75 cents to $1.50 per skin) for wolfers in the Great Plains. Coyote pelts were of significant economic importance during the early 1950s, ranging in price from $5 to $25 per pelt, depending on locality.[204] The coyote's fur is not durable enough to make rugs,[205] but can be used for coats and jackets, scarves, or muffs. The majority of pelts are used for making trimmings, such as coat collars and sleeves for women's clothing. Coyote fur is sometimes dyed black as imitation silver fox.[204]

Coyotes were occasionally eaten by trappers and mountain men during the western expansion. Coyotes sometimes featured in the feasts of the Plains Indians, and coyote pups were eaten by the indigenous people of San Gabriel, California. The taste of coyote meat has been likened to that of the wolf and is more tender than pork when boiled. Coyote fat, when taken in the fall, has been used on occasion to grease leather or eaten as a spread.[206]

Tameability

Coyotes were likely semidomesticated by various pre-Columbian cultures. Some 19th-century writers wrote of coyotes being kept in native villages in the Great Plains. The coyote is easily tamed as a pup, but can become destructive as an adult.[207] Both full-blooded and hybrid coyotes can be playful and confiding with their owners, but are suspicious and shy of strangers,[72] though coyotes being tractable enough to be used for practical purposes like retrieving[208] and pointing have been recorded.[209] A tame coyote named "Butch", caught in the summer of 1945, had a short-lived career in cinema, appearing in Smoky and Ramrod before being shot while raiding a henhouse.[207]

Notes

  1. ^ The name "cased wolf" originates from the fact that the coyote's skin was historically cased like that of the muskrat, whereas the wolf's was spread out flat like the beaver's.[21]
  2. ^ For a full set of supporting references refer to the note (a) in the phylotree at Evolution of the wolf#Wolf-like canids

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  • Olson, Jack (May 2015). The Last Coyote (8 hours). Narrated by Gary MacFadden. Originally published as Slaughter the Animals, Poison the Earth, Simon & Schuster, Oct. 11, 1971. ASIN B00WGUA5HK.

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Coyote: Brief Summary

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The coyote (Canis latrans) is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological niche as the golden jackal does in Eurasia. The coyote is larger and more predatory and was once referred to as the American jackal by a behavioral ecologist. Other historical names for the species include the prairie wolf and the brush wolf.

The coyote is listed as least concern by the International Union for Conservation of Nature, due to its wide distribution and abundance throughout North America. Coyote populations are also abundant southwards through Mexico and into Central America. The species is versatile, able to adapt to and expand into environments modified by humans. It is enlarging its range by moving into urban areas in the eastern U.S. and Canada. The coyote was sighted in eastern Panama (across the Panama Canal from their home range) for the first time in 2013.

The coyote has 19 recognized subspecies. The average male weighs 8 to 20 kg (18 to 44 lb) and the average female 7 to 18 kg (15 to 40 lb). Their fur color is predominantly light gray and red or fulvous interspersed with black and white, though it varies somewhat with geography. It is highly flexible in social organization, living either in a family unit or in loosely knit packs of unrelated individuals. Primarily carnivorous, its diet consists mainly of deer, rabbits, hares, rodents, birds, reptiles, amphibians, fish, and invertebrates, though it may also eat fruits and vegetables on occasion. Its characteristic vocalization is a howl made by solitary individuals. Humans are the coyote's greatest threat, followed by cougars and gray wolves. In spite of this, coyotes sometimes mate with gray, eastern, or red wolves, producing "coywolf" hybrids. In the northeastern regions of North America, the eastern coyote (a larger subspecies, though still smaller than wolves) is the result of various historical and recent matings with various types of wolves. Genetic studies show that most North American wolves contain some level of coyote DNA.

The coyote is a prominent character in Native American folklore, mainly in Aridoamerica, usually depicted as a trickster that alternately assumes the form of an actual coyote or a man. As with other trickster figures, the coyote uses deception and humor to rebel against social conventions. The animal was especially respected in Mesoamerican cosmology as a symbol of military might. After the European colonization of the Americas, it was seen in Anglo-American culture as a cowardly and untrustworthy animal. Unlike wolves, which have undergone an improvement of their public image, attitudes towards the coyote remain largely negative.

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Coywolf

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Captive-bred F1 gray wolf × coyote hybrids, Wildlife Science Center in Forest Lake, Minnesota

Coywolf is an informal term for a canid hybrid descended from coyotes, eastern wolves and gray wolves. All members of the genus Canis are closely genetically related with 78 chromosomes and therefore can interbreed.[1] One genetic study indicates that these two species genetically diverged relatively recently (around 55,000–117,000 years ago). Genomic studies indicate that nearly all North American gray wolf populations possess some degree of admixture with coyotes following a geographic cline, with the lowest levels occurring in Alaska, and the highest in Ontario and Quebec, as well as Atlantic Canada.[2]

Description

Hybrids of any combination tend to be larger than coyotes but smaller than wolves; they show behaviors intermediate between coyotes and the other parent's species.[3][4] In one captive hybrid experiment, six F1 hybrid pups from a male northwestern gray wolf and a female coyote were measured shortly after birth with an average on their weights, total lengths, head lengths, body lengths, hind foot lengths, shoulder circumferences, and head circumferences compared with those on pure coyote pups at birth. Despite being delivered by a female coyote, the hybrid pups at birth were much larger and heavier than regular coyote pups born and measured around the same time.[3] At six months of age, these hybrids were closely monitored at the Wildlife Science Center. Executive Director Peggy Callahan at the facility states that the howls of these hybrids are said to start off much like regular gray wolves with a deep strong vocalization, but changes partway into a coyote-like high pitched yipping.[5]

Compared with pure coyotes, eastern wolf × coyote hybrids form more cooperative social groups and are generally less aggressive with each other while playing.[6] Hybrids also reach sexual maturity when they are two years old, which is much later than occurs in pure coyotes.[7]

Varieties

Eastern coyotes

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Eastern coyote, a coyote-wolf hybrid in West Virginia near the Virginia state line.

Eastern coyotes range from New England, New York, New Jersey, Pennsylvania,[8] Ohio,[9] West Virginia,[10] Maryland,[11] Delaware, and Virginia.[12] Their range also occurs in the Canadian provinces of Ontario, Quebec, New Brunswick,[13] Nova Scotia,[14] Prince Edward Island and Newfoundland and Labrador.[15] Coyotes and wolves hybridized in the Great Lakes region, followed by an eastern coyote expansion, creating the largest mammalian hybrid zone known.[16] Extensive hunting of gray wolves over a period of 400 years caused a population decline that reduced the number of suitable mates, thus facilitating coyote genes swamping into the eastern wolf population. This has caused concern over the purity of remaining wolves in the area, and the resulting eastern coyotes are too small to substitute for pure wolves as apex predators of moose and deer. The main nucleus of pure eastern wolves is currently concentrated within Algonquin Provincial Park. This susceptibility to hybridization led to the eastern wolf being listed as Special Concern under the Canadian Committee on the Status of Endangered Wildlife and with the Committee on the Status of Species at Risk in Ontario. By 2001, protection was extended to eastern wolves occurring on the outskirts of the park, thus no longer depriving Park eastern wolves of future pure-blooded mates. By 2012, the genetic composition of the park's eastern wolves was roughly restored to what it was in the mid-1960s, rather than in the 1980s–1990s, when the majority of wolves had large amounts of coyote DNA.[17]

Aside from the combinations of coyotes and eastern wolves making up most of the modern day eastern coyote's gene pools, some of the coyotes in the northeastern United States also have mild domestic dog (C. lupus familiaris) and western Great Plains gray wolf (C. l. nubilus) influences in their gene pool, thus suggesting that the eastern coyote is actually a four-in-one hybrid of coyotes, eastern wolves, western gray wolves, and dogs, and that the hybrids living in areas with higher white-tailed deer density often have higher degrees of wolf genes than those living in urban environments. The addition of domestic dog genes may have played a minor role in facilitating the eastern hybrids' adaptability to survive in human-developed areas.[18] The four-in-one hybrid theory was further explored in 2014, when Monzón and his team subsequently reanalyzed the tissue and SNP samples taken from 425 eastern coyotes to determine the degree of wolf and dog introgressions involved in each geographic range.[19] The domestic dog allele averages 10% of the eastern coyote's genepool, while 26% is contributed by a cluster of both eastern wolves and western gray wolves. The remaining 64% matched mostly with coyotes. This analysis suggested that prior to the uniformity of its modern-day genetic makeup, multiple swarms of genetic exchanges between the coyotes, feral dogs, and the two distinct wolf populations present in the Great Lakes region may have occurred, and urban environments often favor coyote genes, while the ones in the rural and deep forest areas maintain higher levels of wolf content. A 2016 meta-analysis of 25 genetics studies from 1995 to 2013 found that the northeastern coywolf is 60% western coyote, 30% eastern wolf, and 10% domestic dog. However, this hybrid canid is only now coming into contact with the southern wave of coyote migration into the southern United States.[20]

Red wolves and eastern wolves

The taxonomy of the red and eastern wolf of the Southeastern United States and the Great Lakes regions, respectively, has been long debated, with various schools of thought advocating that they represent either unique species or results of varying degrees of gray wolf × coyote admixture.

In May 2011, an examination of 48,000 single nucleotide polymorphisms in red wolves, eastern wolves, gray wolves, and dogs indicated that the red and eastern wolves were hybrid species, with the red wolf being 76% coyote and only 20% gray wolf, and the eastern wolf being 58% gray wolf and 42% coyote, finding no evidence of being distinct species in either.[21] The study was criticized for having used red wolves with recent coyote ancestry,[22] and a reanalysis in 2012 indicated that it suffered from insufficient sampling.[23] A comprehensive review in 2012 further argued that the study's dog samples were unrepresentative of the species' global diversity, having been limited to boxers and poodles, and that the red wolf samples came from modern rather than historical specimens.[24] The review was itself criticized by a panel of scientists selected for an independent peer review of its findings by the USFWS, which noted that the study's conclusion that the eastern wolf's two unique nonrecombining markers were insufficient to justify full-species status for the animal.[25]

In 2016, a whole-genome DNA study suggested that all of the North American canids, both wolves and coyotes, diverged from a common ancestor 6,000–117,000 years ago. The whole-genome sequence analysis shows that two endemic species of North American wolf, the red wolf and eastern wolf, are admixtures of the coyote and gray wolf.[26][27]

Mexican wolf × coyote hybrids

In a study that analyzed the molecular genetics of coyotes, as well as samples of historical red wolves and Mexican wolves from Texas, a few coyote genetic markers have been found in the historical samples of some isolated Mexican wolf individuals. Likewise, gray wolf Y chromosomes have also been found in a few individual male Texan coyotes.[28] This study suggested that although the Mexican wolf is generally less prone to hybridizations with coyotes, exceptional genetic exchanges with the Texan coyotes may have occurred among individual gray wolves from historical remnants before the population was completely extirpated in Texas. The resulting hybrids would later on melt back into the coyote populations as the wolves disappeared. However, the same study also discussed an alternative possibility that the red wolves, which in turn also once overlapped with both species in central Texas, were involved in circuiting the gene flows between the coyotes and gray wolves, much like how the eastern wolf is suspected to have bridged gene flows between gray wolves and coyotes in the Great Lakes region, since direct hybridizations between coyotes and gray wolves is considered rare.

In tests performed on a stuffed carcass of what was initially labelled a chupacabra, mitochondrial DNA analysis conducted by Texas State University showed that it was a coyote, though subsequent tests revealed that it was a coyote × gray wolf hybrid sired by a male Mexican wolf.[29]

Northwestern wolf × coyote hybrid experiment

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F1 hybrid coyote-gray wolf hybrid, conceived in captivity

In 2013, the U.S. Department of Agriculture Wildlife Services conducted a captive-breeding experiment at their National Wildlife Research Center Predator Research Facility in Logan, Utah. Using gray wolves from British Columbia and western coyotes, they produced six hybrids, making this the first hybridization case between pure coyotes and northwestern wolves. The experiment, which used artificial insemination, was intended to determine whether or not the sperm of the larger gray wolves in the west was capable of fertilizing the egg cells of western coyotes. Aside from the historical hybridizations between coyotes and the smaller Mexican wolves in the south, as well as with eastern wolves and red wolves, gray wolves from the northwestern US and western provinces of Canada were not known to interbreed with coyotes in the wild, thus prompting the experiment.

The six resulting hybrids included four males and two females. At six months of age, the hybrids were closely monitored and were shown to display both physical and behavioral characteristics from both species, as well as some physical similarities to the eastern wolves, whose status as a distinct wolf species or as a genetically distinct subspecies of the gray wolf is controversial. Regardless, the result of this experiment concluded that northwestern wolves, much like the eastern wolves, red wolves, Mexican wolves, and domestic dogs, are capable of hybridizing with coyotes.[3]

In 2015, a research team from the cell and microbiology department of Anoka-Ramsey Community College revealed that an F2 litter of two pups had been produced from two of the original hybrids. At the same time, despite the six F1's successful delivery from the same coyote, they were not all full siblings because multiple sperm from eight different northwestern wolves were used in their production. The successful production of the F2 litter, nonetheless, confirmed that hybrids of coyotes and northwestern wolves are just as fertile as hybrids of coyotes to eastern and red wolves. Both the F1 and F2 hybrids were found to be phenotypically intermediate between the western gray wolves and coyotes. Unlike the F1 hybrids, which were produced via artificial insemination, the F2 litter was produced from a natural breeding.[30]

The study also discovered through sequencing 16S ribosomal RNA encoding genes that the F1 hybrids all have an intestinal microbiome distinct from both parent species, but which was once reported to be present in some gray wolves. Moreover, analysis of their complementary DNA and ribosomal RNA revealed that the hybrids have very differential gene expressions compared to those in gray wolf controls.

Coydogs

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Coydogs in Wyoming

Hybrids between coyotes and domestic dogs have been bred in captivity, which dates to pre-Columbian Mexico.[31] Other specimens were later produced by mammal biologists mostly for research purposes. Domestic dogs are included in the gray wolf species;[32] hence, coydogs are another biological sub-variation of hybrids between coyotes and gray wolves; the dog being considered a domesticated subspecies of Canis lupus.[33]

See also

References

  1. ^ Wayne, R. (1993). "Molecular evolution of the dog family". Trends in Genetics. 9 (6): 218–24. doi:10.1016/0168-9525(93)90122-X. PMID 8337763.
  2. ^ vonHoldt, B. M.; Cahill, J. A.; Fan, Z.; Gronau, I.; Robinson, J.; Pollinger, J. P.; Shapiro, B.; Wall, J.; Wayne, R. K. (2016). "Whole-genome sequence analysis shows that two endemic species of North American wolf are admixtures of the coyote and gray wolf". Science Advances. 2 (7): e1501714. Bibcode:2016SciA....2E1714V. doi:10.1126/sciadv.1501714. PMC 5919777. PMID 29713682.
  3. ^ a b c Mech, L. D.; Christensen, B. W.; Asa, C. S.; Callahan, M.; Young, J. K. (2014). "Production of Hybrids between Western Gray Wolves and Western Coyotes". PLOS ONE. 9 (2): e88861. Bibcode:2014PLoSO...988861M. doi:10.1371/journal.pone.0088861. PMC 3934856. PMID 24586418.
  4. ^ Way J. G. (2007). "A comparison of body mass of Canis latrans (Coyotes) between eastern and western North America" (PDF). Northeastern Naturalist. 14 (1): 111–24. doi:10.1656/1092-6194(2007)14[111:acobmo]2.0.co;2.
  5. ^ Riese, Clive (March 19, 2014), Wildlife Science Center partners in study impacting wolf controversy, Forest Lake Times
  6. ^ Bekoff, M. (1978). "Behavioral Development in Coyotes and Eastern Coyotes", pp. 97–124 in M. Bekoff, (ed.) Coyotes: Biology, Behavior, and Management. Academic Press, New York. ISBN 1930665423.
  7. ^ Way J.G.; Rutledge L.; Wheeldon T.; White B.N. (2010). "Genetic characterization of Eastern "Coyotes" in eastern Massachusetts" (PDF). Northeastern Naturalist. 17 (2): 189–204. doi:10.1656/045.017.0202. S2CID 135542.
  8. ^ "Greater than the sum of its parts". The Economist. October 31, 2015. Retrieved October 30, 2015.
  9. ^ Update on Coy Wolf sightings in Ohio – Ohio Ag Net | Ohio's Country Journal. Ocj.com. Retrieved on 2018-09-05.
  10. ^ West Virginia DNR – Coyote. Wvdnr.gov. Retrieved on 2018-09-05.
  11. ^ Coyotes in Maryland. Department of Natural Resources. maryland.gov
  12. ^ Coyote-Wolf Hybrids Have Spread Across U.S. East. News.nationalgeographic.com (2011-11-08). Retrieved on 2018-09-05.
  13. ^ "Living with Wildlife – Eastern coyotes" (PDF). Natural Resources website. Government of New Brunswick. Retrieved February 2, 2014.
  14. ^ "Frequently Asked Questions about Eastern Coyote in Nova Scotia". Department of Natural Resources website. Government of Nova Scotia. Retrieved February 2, 2014.
  15. ^ "Living with Coyotes in Newfoundland and Labrador". The Department of Environment and Conservation website. Government of Newfoundland and Labrador. Archived from the original on February 19, 2014. Retrieved February 2, 2014.
  16. ^ Vonholdt, Bridgett M.; Kays, Roland; Pollinger, John P.; Wayne, Robert K. (2016). "Admixture mapping identifies introgressed genomic regions in North American canids". Molecular Ecology. 25 (11): 2443–53. doi:10.1111/mec.13667. PMID 27106273.
  17. ^ Rutledge, L. Y.; White, B. N.; Row, J. R.; Patterson, B. R. (2012). "Intense harvesting of eastern wolves facilitated hybridization with coyotes". Ecology and Evolution. 2 (1): 19–33. doi:10.1002/ece3.61. PMC 3297175. PMID 22408723.
  18. ^ Monzón, J; Kays, R; Dykhuizen, D. E. (2013). "Assessment of coyote-wolf-dog admixture using ancestry-informative diagnostic SNPs". Molecular Ecology. 23 (1): 182–197. doi:10.1111/mec.12570. PMC 3899836. PMID 24148003.
  19. ^ Monzon, Javier (January 22, 2014). "It's a "Coyote-wolf-dog eat dog" world". Gotham Coyote Project.
  20. ^ "Northeastern coyote/coywolf taxonomy and admixture: A meta-analysis" (PDF). Canid Biology & Conservation. 19 (1): 1–7. Archived from the original (PDF) on April 4, 2016. Retrieved March 20, 2016.
  21. ^ von Holt, B.M.; et al. (May 12, 2011). "A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids". Genome Res. 21 (8): 1294–305. doi:10.1101/gr.116301.110. PMC 3149496. PMID 21566151.
  22. ^ Beeland, T. DeLene (2013). The Secret World of Red Wolves. Chapel Hill, NC: University of North Carolina Press. ISBN 9781469601991.
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  24. ^ Chambers, Steven M.; Fain, Steven R.; Fazio, Bud; Amaral, Michael (2012). "An account of the taxonomy of North American wolves from morphological and genetic analyses". North American Fauna. 77: 1–67. doi:10.3996/nafa.77.0001.
  25. ^ Dumbacher, J. (January 2014). "Review of Proposed Rule Regarding Status of the Wolf Under the Endangered Species Act" (PDF). NCEAS. U.S. Fish & Wildlife Service.
  26. ^ von Holdt, B.M.; Cahill, J. A.; Fan, Z.; Gronau, I.; Robinson, J.; Pollinger, J.P.; et al. (2016). "Whole-genome sequence analysis shows that two endemic species of North American wolf are admixtures of the coyote and gray wolf". Science Advances. 2 (7): e1501714. Bibcode:2016SciA....2E1714V. doi:10.1126/sciadv.1501714. PMC 5919777. PMID 29713682.
  27. ^ Morell, Virginia (2016). "How do you save a wolf that's not really a wolf?". Science. doi:10.1126/science.aag0699.
  28. ^ Hailer, F.; Leonard, J.A. (2008). "Hybridization among three native North American Canis species in a region of natural sympatry". PLOS ONE. 3 (10): e3333. Bibcode:2008PLoSO...3.3333H. doi:10.1371/journal.pone.0003333. PMC 2556088. PMID 18841199.
  29. ^ Ardizzoni, S. (September 1, 2013). "Texas State University researcher helps unravel mystery of Texas 'blue dog' claimed to be Chupacabra". Bio News Texas.
  30. ^ "NCUR". Archived from the original on April 24, 2016. Retrieved April 1, 2016.
  31. ^ Valadez, Raúl; Rodríguez, Bernardo; Manzanilla, Linda; Tejeda, Samuel. "13. Dog-wolf Hybrid Biotype Reconstruction from the Archaeological City of Teotihuacan in Prehispanic Central Mexico" (PDF). In Snyder, Lynn M.; Moore, Elizabeth A. (eds.). 9th ICAZ Conference, Durham 2002: Dogs and People in Social, Working, Economic, or Symbolic Interaction. pp. 120–130. Archived from the original (PDF) on September 23, 2015. Retrieved December 12, 2014.
  32. ^ "ADW: Canis lupus familiaris: Information". Animal Diversity Web. Retrieved April 1, 2016.
  33. ^ Anderson, T.M.; Vonholdt, B.M.; Candille, S.I.; Musiani, M.; Greco, C.; Stahler, D.R.; et al. (2009). "Molecular and evolutionary history of melanism in North American gray wolves". Science. 323 (5919): 1339–1343. Bibcode:2009Sci...323.1339A. doi:10.1126/science.1165448. PMC 2903542. PMID 19197024.
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Coywolf: Brief Summary

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 src= Captive-bred F1 gray wolf × coyote hybrids, Wildlife Science Center in Forest Lake, Minnesota

Coywolf is an informal term for a canid hybrid descended from coyotes, eastern wolves and gray wolves. All members of the genus Canis are closely genetically related with 78 chromosomes and therefore can interbreed. One genetic study indicates that these two species genetically diverged relatively recently (around 55,000–117,000 years ago). Genomic studies indicate that nearly all North American gray wolf populations possess some degree of admixture with coyotes following a geographic cline, with the lowest levels occurring in Alaska, and the highest in Ontario and Quebec, as well as Atlantic Canada.

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