Coyote

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Coyote
Temporal range: Middle Pleistocene – present (0.74–0.85 Ma) [1] :p131
2009-Coyote-Yosemite.jpg
Mountain coyote (C. l. lestes)
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Canidae
Genus: Canis
Species:
C. latrans
Binomial name
Canis latrans
Say, 1823 [3]
Cypron-Range Canis latrans.svg
Modern range of Canis latrans
Synonyms [4]

The coyote (from Nahuatl coyōtl Loudspeaker.svg pronunciation  ), prairie wolf or brush wolf, Canis latrans, is a canine native to North America. It is smaller than its close relative, the gray wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological niche as the Eurasian golden jackal does in Eurasia, though it is larger and more predatory, and is sometimes called the American jackal by zoologists.

Nahuatl, known historically as Aztec, is a language or group of languages of the Uto-Aztecan language family. Varieties of Nahuatl are spoken by about 1.7 million Nahua peoples, most of whom live in central Mexico.

<i>Canis</i> genus of mammals

Canis is a genus of the Canidae containing multiple extant species, such as wolves, coyotes, jackals, dingoes, and dogs. Species of this genus are distinguished by their moderate to large size, their massive, well-developed skulls and dentition, long legs, and comparatively short ears and tails.

Contents

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, 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, with coyotes moving into urban areas in the Eastern U.S., and was sighted in eastern Panama (across the Panama Canal from their home range) for the first time in 2013.

International Union for Conservation of Nature international organisation

The International Union for Conservation of Nature is an international organization working in the field of nature conservation and sustainable use of natural resources. It is involved in data gathering and analysis, research, field projects, advocacy, and education. IUCN's mission is to "influence, encourage and assist societies throughout the world to conserve nature and to ensure that any use of natural resources is equitable and ecologically sustainable".

Mexico Country in the southern portion of North America

Mexico, officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatemala, Belize, and the Caribbean Sea; and to the east by the Gulf of Mexico. Covering almost 2,000,000 square kilometres (770,000 sq mi), the nation is the fifth largest country in the Americas by total area and the 13th largest independent state in the world. With an estimated population of over 120 million people, the country is the tenth most populous state and the most populous Spanish-speaking state in the world, while being the second most populous nation in Latin America after Brazil. Mexico is a federation comprising 31 states and Mexico City, a special federal entity that is also the capital city and its most populous city. Other metropolises in the state include Guadalajara, Monterrey, Puebla, Toluca, Tijuana and León.

Central America central geographic region of the Americas

Central America is located on the southern tip of North America, or is sometimes defined as a subcontinent of the Americas, bordered by Mexico to the north, Colombia to the southeast, the Caribbean Sea to the east, and the Pacific Ocean to the west and south. Central America consists of seven countries: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, and Panama. The combined population of Central America has been estimated to be 41,739,000 and 42,688,190.

As of 2005, 19 coyote subspecies are recognized. 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. It has a varied diet consisting primarily of animal meat, including 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 United States and eastern Canada, 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.

Subspecies taxonomic rank subordinate to species

In biological classification, the term subspecies refers to one of two or more populations of a species living in different subdivisions of the species' range and varying from one another by morphological characteristics. A single subspecies cannot be recognized independently: a species is either recognized as having no subspecies at all or at least two, including any that are extinct. The term is abbreviated subsp. in botany and bacteriology, ssp. in zoology. The plural is the same as the singular: subspecies.

Fulvous is a colour, sometimes described as dull orange, brownish-yellow or tawny, it can also be likened to a variation of buff, beige or butterscotch. As an adjective it is used in the names of many species of birds, and occasionally other animals, to describe their appearance. It is also used as in mycology to describe fungi with greater colour specificity, specifically the pigmentation of the surface cuticle, the broken flesh and the spores en masse.

Deer A family of mammals belonging to even-toed ungulates

Deer are the hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the fallow deer, and the chital; and the Capreolinae, including the reindeer (caribou), the roe deer, and the moose. Female reindeer, and male deer of all species except the Chinese water deer, grow and shed new antlers each year. In this they differ from permanently horned antelope, which are part of a different family (Bovidae) within the same order of even-toed ungulates (Artiodactyla).

The coyote is a prominent character in Native American folklore, mainly in the Southwestern United States and Mexico, 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 reviled in Anglo-American culture as a cowardly and untrustworthy animal. Unlike wolves (gray, eastern, or red), which have undergone an improvement of their public image, attitudes towards the coyote remain largely negative.

Indigenous peoples of the Americas Pre-Columbian inhabitants of North, Central and South America and their descendants

The indigenous peoples of the Americas are the Pre-Columbian peoples of North, Central and South America and their descendants.

Southwestern United States Geographical region of the USA

The Southwestern United States, also known as the American Southwest, is the informal name for a region of the western United States. Definitions of the region's boundaries vary a great deal and have never been standardized, though many boundaries have been proposed. For example, one definition includes the stretch from the Mojave Desert in California to Carlsbad, New Mexico, and from the Mexico–United States border to the southern areas of Colorado, Utah, and Nevada. The largest metropolitan areas are centered around Phoenix, Las Vegas, Tucson, Albuquerque, and El Paso. Those five metropolitan areas have an estimated total population of more than 9.6 million as of 2017, with nearly 60 percent of them living in the two Arizona cities—Phoenix and Tucson.

Trickster literary archetype

In mythology, and in the study of folklore and religion, a trickster is a character in a story, which exhibits a great degree of intellect or secret knowledge, and uses it to play tricks or otherwise disobey normal rules and conventional behaviour.

Description

Closeup of a Mountain coyote's (C. l. lestes) head Coyote portrait.jpg
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]

Afton, Wyoming Town in Wyoming, United States

Afton is a town in Lincoln County, Wyoming, United States. The population was 1,911 at the 2010 census. Afton is home to the world's largest arch made of elk antlers. Spanning 75 feet (23 m) across the four lanes of US Highway 89, the arch consists of 3,011 elk antlers and weighs 15 tons.

The color and texture of the coyote's fur varies 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]

Desert Area of land where little precipitation occurs

A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to the processes of denudation. About one-third of the land surface of the world is arid or semi-arid. This includes much of the polar regions where little precipitation occurs and which are sometimes called polar deserts or "cold deserts". Deserts can be classified by the amount of precipitation that falls, by the temperature that prevails, by the causes of desertification or by their geographical location.

Albinism Congenital disorder causing skin, eyes, hair/fur, scales, etc. to lack melanin pigmentation

Albinism is the "congenital absence of any pigmentation or coloration in a person, animal or plant, resulting in white hair, feathers, scales and skin and pink eyes in mammals, birds, reptiles, amphibians and fish and other small invertebrates as well." Varied use and interpretation of the terms mean that written reports of albinistic animals can be difficult to verify. Albinism can reduce the survivability of an animal; for example, it has been suggested that albino alligators have an average survival span of only 24 hours due to the lack of protection from UV and their lack of camouflage to avoid predators. Albino animals have characteristic pink or red eyes because the lack of pigment in the iris allows the blood vessels of the retina to be visible. Familiar albino animals include in-bred strains of laboratory animals, but populations of naturally occurring albino animals exist in the wild, e.g. Mexican cave tetra. Albinism is a well-recognized phenomenon in molluscs, both in the shell and in the soft parts. It has been claimed by some, e.g. that "albinism" can occur for a number of reasons aside from inheritance, including genetic mutations, diet, living conditions, age, disease, or injury. However, this is contrary to definitions where the condition is inherited.

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

Toltec pictograph of coyote. Toltec coyote.jpg
Toltec pictograph of 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, distinguishing between coyotes and 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. Lewis, writing on 5 May 1805, in northeastern Montana, described the coyote in these terms:

The small woolf or burrowing dog of the prairies are the inhabitants almost invariably of the open plains; they usually ascociate 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 redish-brown colour. The eye of a deep sea green colour small and piercing. Their [claws] are reather 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, fifteen miles 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) 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] [lower-alpha 1] "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]

Local and indigenous names for Canis latrans

Evolution

Phylogenetic tree of the extant wolf-like canids
Caninae  3.5  Ma
3.0
2.7
1.9
1.6
1.3
1.1
0.8

Domestic dog Tibetan mastiff (white background).jpg

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

Himalayan 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

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

Eurasian golden jackal Dogs, jackals, wolves, and foxes (Plate X).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

Phylogenetic relationships between the extant wolf-like canids based on DNA taken from the cell nucleus, [40] [41] except for the Himalayan wolf (Canis lupus filchneri) that is based on mitochondrial DNA sequences [41] [42] plus X chromosome and Y chromosome sequences. [42] Timing in millions of years. [41]

Fossil record

Skeleton of Pleistocene coyote (C. l. orcutti) Canis latrans orcutti.png
Skeleton of Pleistocene coyote (C. l. orcutti)

Xiaoming Wang and Richard H. Tedford, one of the foremost authorities on carnivore evolution, [43] 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 [44] 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. [45] :p58 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. [1] :p175,180 Johnston describes C. lepophagus as having a more slender skull and skeleton than the modern coyote. [46] :385 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. [47] :p241

C. lepophagus was similar in weight to modern coyotes, but had shorter limb bones that indicates 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 wolf's. 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. [48]

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. [1] :p136 Modern coyotes arose 1,000 years after the Quaternary extinction event. [49] Compared to their modern Holocene counterparts, Pleistocene coyotes (C. l. orcutti) were larger and more robust, likely in response to larger competitors and prey. [49] 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. [50] Their reduction in size occurred within 1000 years of the Quaternary extinction event, when their large prey died out. [49] Furthermore, Pleistocene coyotes were unable to exploit the big-game hunting niche left vacant after the extinction of the dire wolf (C. 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. [50]

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. [51] 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. [52] 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. [53] [54] The proposed timing of the wolf/coyote divergence conflicts with the finding of a coyote-like specimen in strata dated to 1 Mya. [45]

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 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 the Southwestern United States (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

Melanistic coyotes owe their color to a mutation that first arose in domestic dogs. Black coyodog.jpg
Melanistic coyotes owe their color to a mutation that first arose in domestic dogs.

Coyotes have occasionally mated with 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]

Coywolf hybrid conceived in captivity between a male gray wolf and a female coyote Westerncoywolf.png
Coywolf hybrid conceived in captivity between a male gray wolf and a female coyote

Coyotes have hybridized with wolves to varying degrees, particularly in the Eastern United States and Canada. 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 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 United States, 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. [53] [80]

Behavior

Social and reproductive behaviors

Mearns' coyote (C. l. mearnsi) pups playing Gpa bill coyote pups 3.jpg
Mearns' coyote (C. l. mearnsi) pups playing
A pack of coyotes in Yellowstone National Park Pack of coyotes on snow.jpg
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]

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]

Communication

A coyote howling Howl (cropped).jpg
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. [101] 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. [102] 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. [101]

Vocalizations

The coyote has been described as "the most vocal of all [wild] North American mammals". [103] [104] 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 as 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

A Sonoran Desert coyote at the Sonora Desert Museum in Tucson Arizona Coyote at Sonora Desert Museum Tucson Arizona.jpg
A Sonoran Desert coyote at the Sonora Desert Museum in Tucson Arizona
Urban coyote in Bernal Heights, San Francisco Urban Coyote, Bernal Heights.jpg
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. [105]

Diet

Coyote with scrap of road killed pronghorn in Seedskadee National Wildlife Refuge, Wyoming Coyote at Seedskadee National Wildlife Refuge (31034864347).jpg
Coyote with scrap of road killed pronghorn in Seedskadee National Wildlife Refuge, Wyoming

The coyote is roughly the North American equivalent to the Eurasian golden jackal. [106] 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, deer, sheep, rabbits, rodents, birds, amphibians (except toads), lizards, snakes, 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] More unusual prey include fishers, [107] young black bear cubs, [108] harp seals [109] 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. [110] Birds taken by coyotes may range in size from thrashers, larks and sparrows to adult wild turkeys and, possibly, brooding adult swans and pelicans. [111] [112] [113] [114] 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). [115] [116] 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 calves. 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. [115] [117] [118] [119] [120] [121] [122]

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. [123] In Death Valley, coyotes may consume great quantities of hawkmoth caterpillars or beetles in the spring flowering months. [124]

Enemies and competitors

Comparative illustration of coyote and gray wolf USFWS - How to recognise a gray wolf.png
Comparative illustration of coyote and gray wolf
Mountain coyotes (C. l. lestes) cornering a juvenile cougar Feeling Unwelcome.jpg
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. [125] [126] 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 Nevadas, 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. [127] Coyotes that are killed are sometimes not eaten, perhaps indicating that these compromise competitive interspecies interactions, however there are multiple confirmed cases of cougars also eating coyotes. [128] [129] 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. [130]

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, [131] American alligators, [132] large Canada lynx [133] and golden eagles. [134] 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. [135] [136] [137] [138] [139] [140] [141] [142]

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. [143] In southern California, coyotes frequently kill gray foxes, and these smaller canids tend to avoid areas with high coyote densities. [144]

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. [145] However, several studies have demonstrated interference competition between coyotes and bobcats, and in all cases coyotes dominated the interaction. [146] [147] Multiple researchers [148] [149] [150] [147] [151] reported instances of coyotes killing bobcats, whereas bobcats killing coyotes is more rare. [146] 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. [147] 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. [144] 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 Canadian lynx on occasion, [152] [153] [154] and compete with them for prey, especially snowshoe hares. [152] 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. [155]

Range

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) Tiburon 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 Coyote subspecies distribution map.svg
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
Coyote expansion over the past 10,000 years Coyote expansion past 10,000 years.jpg
Coyote expansion over the past 10,000 years
Coyote expansion over the decades since 1900 Coyote expansion by decade.jpg
Coyote expansion over the decades since 1900

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 the United States and Canada, 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 all of the United States and Mexico, southward into Central America, and northward into most of Canada and Alaska. [157] 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 rainforests. The coyote is predicted to appear in northern Belize in the near future, as the habitat there is favorable to the species. [158] 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. [159] 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 recent genetic study proposes that coyotes were originally not found in 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. [160]

Diseases and parasites

California valley coyote (C. l. ochropus) suffering from sarcoptic mange Mangy coyote Ano Nuevo State Park.jpg
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. [161] 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 through their rodent and lagomorph prey, and can be deadly for pups. [162]

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. [163] 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. [162]

Relationships with humans

In folklore and mythology

Kojote2010 cut.JPG

Coyote features as a trickster figure and skin-walker in the folktales of some Native Americans in the United States, 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. [164] Folklorists such as Harris believe coyotes came to be seen as tricksters due to the animal's intelligence and adaptability. [165] After the European colonization of the Americas, Anglo-American depictions of Coyote are of a cowardly and untrustworthy animal. [166] Unlike the gray wolf, which has undergone a radical improvement of its public image, Anglo-American cultural attitudes towards the coyote remain largely negative. [167]

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. [168]

Mural from Atetelco, Teotihuacan depicting coyote warriors. Teotihuacan - Palacio de Atetelco Wandmalerei 3.jpg
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. [169] 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. [170] He is sometimes depicted as a womanizer, responsible for bringing war into the world by seducing Xochiquetzal, the goddess of love. [171] 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. [172]

Attacks on humans

A sign discouraging people from feeding coyotes, which can lead to them habituating themselves to human presence, thus increasing the likelihood of attacks No Feeding.jpg
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 [173] and another on a nineteen-year-old named Taylor Mitchell in Nova Scotia, Canada. [174] 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. [175] 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. [173]

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. [173] Non-rabid coyotes in these areas sometimes target small children, mostly under the age of 10, though some adults have been bitten. [176]

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. [177]

Livestock and pet predation

Coyote confronting a dog Coyote vs Dog.jpg
Coyote confronting a dog

Coyotes are presently the most abundant livestock predators in western North America, causing the majority of sheep, goat, and cattle losses. [178] 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. [179] The total number of sheep deaths in 2004 comprised 2.22% of the total sheep and lamb population in the United States, [180] 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. [181] 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. [182] An Idaho census taken in 2005 showed that individual coyotes were 5% as likely to attack livestock as individual wolves. [183] In Utah, more than 11,000 coyotes were killed for bounties totaling over $500,000 in the fiscal year ending June 30, 2017. [184]

Livestock guardian dogs are commonly used to aggressively repel predators and have worked well in both fenced pasture and range operations. [185] A 1986 survey of sheep producers in the USA found that 82% reported the use of dogs represented an economic asset. [186]

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. [187] 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. [188]

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. [178]

Coyote with a typical throat hold on domestic sheep Coyote with typical hold on lamb.jpg
Coyote with a typical throat hold on domestic sheep

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. [178] 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. [189] [190]

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. [191] Scat analysis collected near Claremont, California, revealed that coyotes relied heavily on pets as a food source in winter and spring. [173] 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. [173] Coyotes usually attack smaller-sized dogs, but they have been known to attack even large, powerful breeds such as the Rottweiler in exceptional cases. [192] Dogs larger than coyotes, such as greyhounds, are generally able to drive them off, and have been known to kill coyotes. [193] Smaller breeds are more likely to suffer injury or death. [176]

Uses

Fur of a Canadian coyote Canis latrans (Kanada) fur skin.jpg
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. [194] The coyote's fur is not durable enough to make rugs, [195] 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. [194]

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. [196]

Tameability

Coyotes were probably 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. [197] 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 [198] and pointing have been recorded. [199] 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. [197]

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]

Related Research Articles

Canidae family of mammals

The biological family Canidae is a lineage of carnivorans that includes domestic dogs, wolves, coyotes, foxes, jackals, dingoes, and many other extant and extinct dog-like mammals. A member of this family is called a canid.

Jackal several species of the wolf genus of mammals

Jackals are medium-sized omnivorous mammals of the genus Canis, which also includes wolves, coyotes and the domestic dog. While the word "jackal" has historically been used for many small canids, in modern use it most commonly refers to three species: the closely related black-backed jackal and side-striped jackal of sub-Saharan Africa, and the golden jackal of south-central Eurasia, which is more closely related to other members of the genus Canis.

Red wolf subspecies of mammal

The red wolf is a canine native to the southeastern United States. The subspecies is the product of ancient genetic admixture between the gray wolf and the coyote, however it is regarded as unique and therefore worthy of conservation. Morphologically it is intermediate between the coyote and gray wolf, and is of a reddish, tawny color. The US Endangered Species Act of 1973 currently does not provide protection for endangered admixed individuals and researchers argue that these should warrant full protection under the Act. However, the red wolf when considered as a species is listed as an endangered species under this Act and is protected by law. Although Canis rufus is not listed in the CITES Appendices of endangered species, since 1996 the IUCN has listed it as a critically endangered species.

Wolf species of mammal

The wolf, also known as the gray/grey wolf, timber wolf, or tundra wolf, is a canine native to the wilderness and remote areas of Eurasia and North America. It is the largest extant member of its family, with males averaging 43–45 kg (95–99 lb) and females 36–38.5 kg (79–85 lb). It is distinguished from other Canis species by its larger size and less pointed features, particularly on the ears and muzzle. Its winter fur is long and bushy and predominantly a mottled gray in color, although nearly pure white, red and brown to black also occur. Mammal Species of the World, a standard reference work in zoology, recognises 38 subspecies of C. lupus.

Maned wolf species of mammal

The maned wolf is the largest canid of South America. Its markings resemble those of foxes, but it is neither a fox nor a wolf. It is the only species in the genus Chrysocyon.

Wolfdog Dog breed

A wolfdog is a canine produced by the mating of a domestic dog with a gray wolf, eastern timber wolf, red wolf, or Ethiopian wolf to produce a hybrid.

Eastern wolf subspecies of mammal

The eastern wolf is a subspecies of gray wolf native to the Great Lakes region and southeastern Canada. The subspecies is the product of ancient genetic admixture between the gray wolf and the coyote, however it is regarded as unique and therefore worthy of conservation. There are two forms, the larger being referred to as the Great Lakes wolf and the smaller being the Algonquin wolf. The eastern wolf's morphology is midway between that of the northwestern wolf and the coyote. The fur is typically of a grizzled grayish-brown color mixed with cinnamon. The nape, shoulder and tail region are a mix of black and gray, with the flanks and chest being rufous or creamy. It primarily preys on white-tailed deer, but may occasionally attack moose and beavers.

Ethiopian wolf species of mammal

The Ethiopian wolf, also known as the Simien jackal or Abyssinian wolf, is a canid native to the Ethiopian Highlands. It is similar to the coyote in size and build, and is distinguished by its long and narrow skull, and its red and white fur. Unlike most large canids, which are widespread, generalist feeders, the Ethiopian wolf is a highly specialised feeder of Afroalpine rodents with very specific habitat requirements. It is one of the world's rarest canids, and Africa's most endangered carnivore.

Golden jackal A wolf-like canid that is native to Southeast Europe and Asia

The golden jackal is a wolf-like canid that is native to Southeast Europe, Southwest Asia, South Asia, and regions of Southeast Asia. Compared with the Arabian wolf, which is the smallest of the gray wolves, the jackal is smaller and possesses shorter legs, a shorter tail, a more elongated torso, a less-prominent forehead, and a narrower and more pointed muzzle. The golden jackal's coat can vary in color from a pale creamy yellow in summer to a dark tawny beige in winter. It is listed as Least Concern on the IUCN Red List due to its widespread distribution and high density in areas with plenty of available food and optimum shelter.

Coydog hybrid mammal

A coydog is a canid hybrid resulting from a mating between a coyote and a dog. The term is sometimes mistakenly used for coywolves, which are common in northeast North America, whereas true coydogs are only occasionally found in the wild. A study found that when a coyote met a dog, the reaction was either antagonistic or equally as likely to lead to bouts of play.

Coywolf hybrid mammal

Coywolf is an informal term for a canid hybrid descended from coyotes, eastern wolves and gray wolves. All members of the genus Canis are genetically closely related because their chromosomes number 78, therefore they can interbreed. One genetic study indicates that these two species genetically diverged relatively recently. 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.

Mexican wolf subspecies of mammal

The Mexican wolf, also known as the lobo, is a subspecies of gray wolf once native to southeastern Arizona, southern New Mexico, western Texas and northern Mexico. It is the smallest of North America's gray wolves, and is similar to C. l. nubilus, though it is distinguished by its smaller, narrower skull and its darker pelt, which is yellowish-gray and heavily clouded with black over the back and tail. Its ancestors were likely the first gray wolves to enter North America after the extinction of the Beringian wolf, as indicated by its southern range and basal physical and genetic characteristics.

Canid hybrids are the result of interbreeding between different species of the canine (dog) family. They often occur in the wild, in particular between domestic or feral dogs and wild native canids.

Indian wolf subspecies of mammal

The Indian wolf is a subspecies of grey wolf that ranges from Southwest Asia to the Indian Subcontinent. It is intermediate in size between the Tibetan and Arabian wolf, and lacks the former's luxuriant winter coat due to it living in warmer conditions. Two closely related haplotypes within this subspecies have been found basal to all other extant Canis lupus haplotypes apart from the older-lineage Himalayan wolf, and have been proposed as a separate species.

<i>Canis lepophagus</i> species of mammal

Canis lepophagus is an extinct species of canid which was endemic to much of North America during the Early Pliocene. It is notable because its lineage is proposed to have led to both wolves and coyotes.

Eastern coyote canine hybride

The eastern coyote is a wild North American canine of mixed coyote-wolf and dog parentage. The hybridization likely first occurred in the Great Lakes region, as western coyotes moved east. It was first noticed during the early 1930s to the late 1940s, and likely originated in the aftermath of the extirpation of the gray wolf in southeastern Ontario, Labrador and Quebec, thus allowing coyotes to colonize the former wolf ranges and mix with the remnant wolf populations. This hybrid is smaller than the eastern wolf and holds smaller territories, but is larger and holds more extensive home ranges than the typical western coyote.

Pleistocene coyote subspecies of mammal

The Pleistocene coyote, also known as the Ice Age coyote, is an extinct subspecies of coyote that lived in western North America during the Late Pleistocene era. Most remains of the subspecies were found in southern California, though at least one was discovered in Idaho. It was part of a carnivore guild that included other canids like foxes, gray wolves, and dire wolves.

African golden wolf виды млекопитающих

The African golden wolf, also known as the Egyptian jackal or grey jackal, is a canid native to north and northeastern Africa. The species is the descendant of a genetically admixed canid of 72% grey wolf and 28% Ethiopian wolf ancestry. The species is common in northwest and northeast Africa, occurring from Senegal to Egypt in the east, in a range including Morocco, Algeria, Tunisia and Libya in the north to Nigeria, Chad and Tanzania in the south. It is a desert-adapted canid, and is common in plains and steppe areas, including ones lacking abundant water. In the Atlas Mountains, the species has been sighted in elevations as high as 1,800 metres. It is primarily a predator, targeting invertebrates and mammals as large as gazelle fawns, though larger animals are sometimes taken. Other foodstuffs include animal carcasses, human refuse, and fruit. The African golden wolf is a monogamous and territorial animal, whose social structure includes yearling offspring remaining with the family to assist in raising their parents' younger pups.

Evolution of the wolf

The evolution of the wolf occurred over a geologic time scale of at least 300 thousand years. The grey wolf Canis lupus is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the Holarctic. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.

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