Coyote

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For other uses, see Coyote (disambiguation).
Coyote
Temporal range: 0.7–0 Ma
Middle Pleistocene – recent
2009-Coyote-Yosemite.jpg
Canis latrans lestes
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Class: Mammalia
Order: Carnivora
Suborder: Caniformia
Family: Canidae
Genus: Canis
Species: C. latrans
Binomial name
Canis latrans
Say, 1823
Cypron-Range Canis latrans.svg
Modern range of Canis latrans

The coyote (US /kˈti, ˈkt/, UK /kɔɪˈjt, kɔɪˈjt/;[a] Canis latrans) is a canid native to North America and Central America. It is smaller than its close relative, the gray wolf, being roughly the North American equivalent to the Old World golden jackal, though it is larger and more predatory. It is listed as "least concern" by the International Union for Conservation of Nature (IUCN), on account of its wide distribution and abundance throughout North America, southwards through Mexico and into Central America. The species is versatile and able to adapt to environments modified by humans. As human activity has altered the landscape, the coyote's range has expanded. In 2013, coyotes were sighted in eastern Panama (across the Panama Canal from their home range) for the first time. The coyote is more closely related to the common ancestor of wolves and other canids (more "basal") than its relative the gray wolf. As of 2005, 19 coyote subspecies are recognized.

The ancestors of the coyote diverged from those of gray wolves 6,000–117,000 years ago, with the modern species arising in North America after the Quaternary extinction event. 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. The coyote's characteristic vocalization is a howl made by solitary individuals. Humans aside, cougars and gray wolves are the coyote's only serious enemies. Nevertheless, coyotes have on occasion mated with the gray wolf, producing hybrids colloquially called "coywolves".

The coyote is a prominent character in Native American folklore (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 acts as a picaresque hero which rebels against social convention through deception and humor. The animal was especially respected in Mesoamerican cosmology as a symbol of military might, with some scholars having traced the origin of the Aztec god Quetzalcoatl to a pre-Aztec coyote deity. After the European colonization of the Americas, it was reviled in Anglo-American culture as a cowardly and untrustworthy animal. Unlike the gray wolf, which has undergone a radical improvement of its public image, cultural attitudes towards the coyote remain largely negative.

Description[edit]

Life study by Ernest Thompson Seton (1886).

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 to 1.5 m (3 ft 3 in to 4 ft 11 in), and tail length 40 cm (16 in), with females being shorter in both body length and height.[2] 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).[3] Scent glands are located at the upper side of the base of the tail and are a bluish-black color.[4]

The color and texture of the coyote's fur varies somewhat geographically.[2] 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.[5] 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).[6] Albinism is extremely rare in coyotes; out of a total of 750,000 coyotes harvested by federal and cooperative hunters between March 22, 1938, and June 30, 1945, only two were albinos.[5]

The coyote is typically smaller than the gray wolf, but has longer ears and a larger braincase,[2] as well as a thinner frame, face, and muzzle. The scent glands are smaller than the gray wolf's, but are the same color.[4] Its fur color variation is much less varied than that of a wolf.[7] The coyote also carries its tail downwards when running or walking, rather than horizontally as the wolf does.[8]

Coyote tracks can be distinguished from those of dogs by their more elongated, less rounded shape.[9][10]

Taxonomy and evolution[edit]

History[edit]

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.[11] It is often difficult in early post-Columbian historical records to distinguish between coyotes and wolves. 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.[12] 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 May 5, 1805, in northeastern Montana, described the coyote in the following terms:[13]

Toltec pictograph of coyote.

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 tallons [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.

The coyote was first scientifically described by 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 May 5, 1805.[14][15]

Naming and etymology[edit]

The earliest written reference to the species comes from 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" (the root word of which is the Nahuatl coyōtl meaning "barking dog") comes from Francisco Javier Clavijero's Historia de México in 1780.[16] 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.[13][17] Alternative English names for the coyote include "prairie wolf", "brush wolf", "cased wolf",[18][b] "little wolf"[19] and "American jackal".[20]

Local and indigenous names for Canis lantrans

Evolution[edit]

Skeleton of Pleistocene coyote (C. l. orcutti)

The earliest fossil carnivores that can be linked with some certainty to canids are the Eocene miacids, which lived some 38 to 56 million years ago (mya). The miacids later diverged into the feliforms and the caniforms, with the former line leading to such genera as the coyote-sized Mesocyon of the Oligocene (38 to 24 mya), the fox-like Leptocyon, and the wolf-like Tomarctus which inhabited North America some 10 mya.[36] 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.[37] Modern phylogenetics places the coyote between the gray wolf and golden jackal.[38] Coyotes and gray wolves diverged 6,000 to 117,000 years ago in the Late Pleistocene to Middle Holocene periods.[39]





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



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












Dog Dogs, jackals, wolves, and foxes (Plate XXXVII).jpg



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




Coyote Dogs, jackals, wolves, and foxes (Plate IX).jpg[38]




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





The evolution of the coyotes is remarkably well documented, and can be traced back in an unbroken line to the Hemphillian Eucyon davisi.[40] The coyote likely arose from a certain population of the Blancan species C. lepophagus which,[41] although similar in weight to modern coyotes, had shorter limb bones, thus indicating a less cursorial lifestyle.[37] Modern variety arose 1,000 years after the Quaternary extinction event.[42] Compared to their modern Holocene counterparts, Pleistocene coyotes (C. l. orcutti) were larger and more robust, likely in response to larger competitors and prey.[42] 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.[43] Their reduction in size occurred within 1000 years of the Quaternary extinction event, when their large prey died out.[42] Furthermore, Pleistocene coyotes were unable to exploit the big-game hunting niche left vacant after the extinction of the dire wolf, as it was rapidly filled by gray wolves, which likely actively killed off the large coyotes, with natural selection favoring the modern gracile morph.[43]

Subspecies[edit]

As of 2005, 19 subspecies are recognized.[44] 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 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.[45]

Hybridization[edit]

Melanistic coyote

Coyotes have occasionally mated with dogs, sometimes producing crosses colloquially known as "coydogs".[57] 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, with even captive specimens having shown reluctance to mate with them. 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.[58] In captivity, F1 hybrids tend to be more mischievous and less manageable as pups than dogs, and are less trustworthy on maturity than wolf-dog hybrids.[57] Hybrids vary in appearance, but generally retain the coyote's adult sable coat color, dark neonatal coat color, bushy tail with an active supracaudal gland, and white facial mask. F1 hybrids tend to be intermediate in form between dogs and coyotes, while F2 hybrids are more varied. Both F1 and F2 hybrids resemble their coyote parents in terms of shyness and intrasexual aggression.[59] Hybrid play behavior includes the coyote "hip-slam".[60] Hybrids of both sexes are fertile, and can be successfully bred through four generations.[57] Melanistic coyotes owe their black pelts to a mutation that first arose in domestic dogs.[61] A population of nonalbino white coyotes in Newfoundland owe their coloration to a melanocortin 1 receptor mutation inherited from Golden Retrievers.[62]

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 has been confirmed to be of mixed wolf-coyote parentage, and probably originated in the aftermath of the extermination of wolves in the northeast, thus allowing coyotes to colonize former wolf ranges and mix with remnant wolf populations. This hybrid is smaller than the 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.[63] Adult eastern coyotes are larger than western coyotes, weighing an average of 14 to 18 kg (30 to 40 lb), with female eastern coyotes weighing 21% more than male western coyotes.[63][64] Eastern coyotes also weigh more at birth; while newborn western coyotes weigh 250 to 300 g (0.6 to 0.7 lb), eastern coyotes weigh 350 to 360 g (0.77 to 0.79 lb). By the age of 35 days, eastern coyote pups average 1.6 kg (4 lb), which is 200 g (0.4 lb) more than western coyotes of similar age. By this time, physical differences become more apparent, 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.[65]

Aside from its size, the eastern coyote is physically similar to the western coyote; both have erect ears, a straight and bushy tail, a conspicuous supracaudal gland, and a narrow chest. 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.[66] No significant differences are seen between eastern and western coyotes in expressing 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.[65] Eastern coyotes tend to reach sexual maturity at two years of age, much later than in western coyotes.[63] In 2011, an analysis of 48,000 single-nucleotide polymorphism chips in the genomes of various wolf and coyote populations revealed that the eastern wolf (native to Algonquin Provincial Park) and the red wolf (native to North Carolina), both previously labeled as species distinct from the gray wolf, are in fact products of varying degrees of wolf-coyote hybridization. The wolf-coyote admixture resulting in the development of the eastern wolf may have occurred on the order of 600–900 years ago between gray wolves and a now-extinct pre-Columbian coyote population. The eastern wolf has since backcrossed extensively with parent gray wolf populations. The red wolf may have originated later, about 287–430 years ago, when much of the southeastern United States was being converted to agriculture and predators were targeted for extermination. During this period, declining local wolf populations would have been forced to mate with coyotes, with the resulting hybrids backcrossing to coyotes as the wolves disappeared, to the extent that about 75–80% of the modern red wolf's genome is of coyote derivation.[67] These findings were corroborated in a further genomic study conducted in 2016.[39]

Behavior[edit]

Social and reproductive behaviors[edit]

Mearns' coyote (C. l. mearnsi) pups playing
A pack of coyotes

Like the 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.[68] 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.[19] Pair bonding can occur 2–3 months before actual copulation takes place.[69] A female entering estrus attracts males by scent marking[70] and howling with increasing frequency.[20] 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.[19] Unlike the wolf, which has been known to practice both monogamous and bigamous matings,[71] the coyote is strictly monogamous, even in areas with high coyote densities and abundant food.[72] 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.[19] 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.[20]

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.[19] Unlike wolf pups, coyote pups begin seriously fighting prior to engaging in play behavior. 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.[73] 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.[19]

Territorial and sheltering behaviors[edit]

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,[19] and is much less aggressive towards intruders than the wolf is, typically chasing and sparring with them, but rarely killing them.[74] Conflicts between coyotes can arise during times of food shortage.[19]

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.[75] A single den can be used year after year.[20]

Hunting and feeding behaviors[edit]

While the popular consensus is that olfaction is very important for hunting,[76] 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[77] and coyotes.[78][79]

When hunting large prey, the coyote often works in pairs or small groups.[2] 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.[20] 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.[80] Coyotes catch mouse-like rodents by pouncing, whereas ground squirrels are chased. Although coyotes can live in large groups, small prey is typically caught singly.[20] 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.[81] Coyotes sometimes urinate on their food, possibly to claim ownership over it.[82]

Coyotes may occasionally form mutualistic relationships with American badgers, assisting each other in digging up rodent prey. 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.[83]

Ecology[edit]

Habitat[edit]

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, antelope, 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.[19]

Coyotes walk around 5-16 km 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 0.8km across water.[84]

Diet[edit]

The coyote is highly versatile in its choice of food, but is primarily carnivorous, with 90% of its diet consisting of animal 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.[19] More unusual prey include fishers,[85] young black bear cubs,[86] harp seals[87] 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.[88] In Death Valley, coyotes may consume great quantities of hawkmoth caterpillars or beetles in the spring flowering months.[89] 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.[19] The coyote readily cannibalizes the carcasses of conspecifics, with coyote fat having been successfully used by coyote hunters as a lure or poisoned bait.[4] 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.[2]

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 corn, wheat, and sorghum.[19]

Enemies and competitors[edit]

Comparative illustration of coyote and gray wolf
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.[90][91] Wolves have been observed to not tolerate coyotes in their vicinity, though coyotes have been known to trail wolves to feed on their kills.[83]

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.[92] In southern California, coyotes frequently kill gray foxes, and these smaller canids tend to avoid areas with high coyote densities.[93]

Coyotes may compete with cougars in some areas. In the eastern Sierra Nevadas, coyotes compete with cougars over mule deer. Cougars usually outcompete coyotes, and may kill them occasionally, thus reducing coyote predation pressure on smaller carnivores such as foxes and bobcats.[94]

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.[95] However, several studies have demonstrated interference competition between coyotes and bobcats, and in all cases coyotes dominated the interaction.[96][97] Instances of bobcats killing coyotes are rarer than instances of coyotes killing bobcats.[96][98] 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.[97] 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.[93] 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.[83] Coyotes have been documented to directly kill Canadian lynx on occasion,[99][100][101] and compete with them for prey, especially snowshoe hares.[99] 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.[102]

Range[edit]

Range of coyote subspecies as of 1978, (1) Mexican coyote, (2) San Pedro Martir coyote, (3) 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[47]

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. This expansion is ongoing, and the species now occupies the majority of areas between 8°N (Panama) and 70°N (northern Alaska).[1]

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.[103] 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.[104] This fear was partially confirmed in January 2013, when the species was recorded in eastern Panama's Chepo District, beyond the Panama Canal.[50]

Communication[edit]

Body language[edit]

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.[68] 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.[105] 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.[106] 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.[105]

Vocalizations[edit]

The coyote has been described as "the most vocal of all [wild] North American mammals".[107][108] 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.[109]

Diseases and parasites[edit]

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.[110] 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.[111]

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 areas 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.[112] Tapeworms have been recorded to infest 60–95% of all coyotes examined. The most common species to infest coyotes is Taenia pisiformis and T. 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). Though 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.[111]

Relationships with humans[edit]

Further information: Urban coyote

The coyote is a prominent character in Native American folklore (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 acts as a picaresque hero which rebels against social convention through deception and humor.[113] The animal was especially respected in Mesoamerican cosmology as a symbol of military might,[114] with some scholars having traced the origin of the Aztec god Quetzalcoatl to a pre-Aztec coyote deity.[115] After the European colonization of the Americas, it was reviled in Anglo-American culture as a cowardly and untrustworthy animal.[116] Unlike the gray wolf, which has undergone a radical improvement of its public image, cultural attitudes towards the coyote remain largely negative.[117]

In folklore and mythology[edit]

Main article: Coyote (mythology)
Spirit shield fashioned from coyote skull and crow feathers

The coyote features prominently as a trickster figure in the folktales of America's indigenous peoples, alternately assuming the form of an actual coyote or a man. As with other trickster figures, the coyote acts as a picaresque hero which rebels against social convention through deception and humor.[113] The coyote was likely given its trickster role in light of the actual animal's intelligence and adaptability; pre-Columbian American people observed its behavior, and their folkloric representations reflected its attributes.[118] It is variously credited for having brought fire to humanity, releasing the bison into the world, and of having slain monsters by petrifying them. The Maidu creation myth has the coyote introducing work, suffering, and death to the world. Zuni folklore has the coyote bringing winter into the world by stealing light from the kachinas. Some tribes, such as the Chinook, Maidu, Paiute, Pawnee, Tohono O'odham, and Ute portray the coyote as the companion of the creator. In the Paiute creation myth, the coyote was created by the wolf as a companion, and the two created land by piling soil on the water-covered world. A Tohono O'odham flood myth has the coyote helping Montezuma survive a global deluge that destroys humanity. After the Great Mystery creates humanity, the coyote and Montezuma teach people how to live. The Crow creation myth portrays Old Man Coyote as the creator. In Navajo mythology, the 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.[119]

Prior to the Spanish conquest of the Aztec Empire, the coyote played a significant role in Mesoamerican cosmology. The coyote symbolized military might in pre-Aztec 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 Aztec rule.[114] 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.[120] He is sometimes depicted as a womanizer, responsible for bringing war into the world by seducing Xochiquetzal, the goddess of love.[121] 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.[115]

Attacks on humans[edit]

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

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. 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.[122] 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.[123]

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.[123] Non-rabid coyotes in these areas sometimes target small children, mostly under the age of 10, though some adults have been bitten.[124]

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.[125]

Livestock and pet predation[edit]

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.[126] 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.[127] The total number of sheep deaths in 2004 comprised 2.22% of the total sheep and lamb population in the United States.[128] According to the National Agricultural Statistics Service USDA report, "All sheep and lamb inventory in the United States on July 1, 2005, totaled 7.80 million head, 2% above July 1, 2004. Breeding sheep inventory at 4.66 million head on July 1, 2005 was 2% above July 1, 2004."[129] Because coyote populations are typically many times greater and more widely distributed than those of wolves, coyotes cause more overall predation losses. The United States government agents routinely shoot, poison, trap, and kill about 90,000 coyotes each year to protect livestock.[130] An Idaho census taken in 2005 showed that individual coyotes were 5% as likely to attack livestock than individual wolves.[131]

Livestock guardian dogs are commonly used to aggressively repel predators and have worked well in both fenced pasture and range operations.[132] A 1986 survey of sheep producers in the USA found that 82% reported the use of dogs represented an economic asset.[133]

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.[126]

Coyote with a typical throat hold on domestic sheep

Coyote predation can usually be distinguished from dog or coydog predation because coyotes partially consume their victims. Tracks are also 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.[126] 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.[134]

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.[135] Scat analysis collected near Claremont, California, revealed that coyotes relied heavily on pets as a food source in winter and spring.[123] 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.[123] Coyotes usually attack smaller-sized dogs, but they have been known to attack even large, powerful breeds such as the Rottweiler in exceptional cases.[136] Dogs larger than coyotes, such as greyhounds, are generally able to drive them off, and have been known to kill coyotes.[137] Smaller breeds are more likely to suffer injury or death.[124]

Uses[edit]

Prior to the mid-1800s, 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.[138] The coyote's fur is not durable enough to make rugs,[139] 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.[138]

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.[140]

Tameability[edit]

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.[141] Both full-blooded and hybrid coyotes can be playful and confiding with their owners, but are suspicious and shy of strangers,[57] though coyotes being tractable enough to be used for practical purposes like retrieving[142] and pointing have been recorded.[143] 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.[141]

Notes[edit]

  1. ^ Respelled US ky-OHT-ee, KY-oht and UK koy-YOHT-ay, koy-YOHT
  2. ^ 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.[18]

References[edit]

  1. ^ a b Sillero-Zubiri & Hoffmann (2008). Canis latrans. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved May 5, 2008.
  2. ^ a b c d e Bekoff M. (1977). "Canis latrans" (PDF). Mammalian Species. 79 (79): 1–9. doi:10.2307/3503817. ISSN 1545-1410. JSTOR 3503817. OCLC 46381503. 
  3. ^ Young & Jackson 1978, p. 48
  4. ^ a b c Young & Jackson 1978, pp. 63–4
  5. ^ a b Young & Jackson 1978, pp. 50–53
  6. ^ Young & Jackson 1978, p. 247
  7. ^ "Sharing the Land with Wolves" (PDF). Wisconsin Department of Natural Resources. 2015. Retrieved June 29, 2016. 
  8. ^ Cartaino 2011, p. 16
  9. ^ Young & Jackson 1978, p. 59
  10. ^ Vantassel, Stephen (2012). "Coyotes". Wildlife Damage Inspection Handbook (3 ed.). Lincoln, Nebraska: Wildlife Control Consultant. p. 112. ISBN 978-0-9668582-5-9. OCLC 794471798. 
  11. ^ Nowak 1979, p. 14
  12. ^ Hoffmeister, Donald F. (2002). Mammals of Illinois. University of Illinois Press. pp. 33–34. ISBN 978-0-252-07083-9. OCLC 50649299. 
  13. ^ a b Mussulman, Joseph (November 2004). "Coyote". Discovering Lewis & Clark. Retrieved January 15, 2013. 
  14. ^ James, Edwin; Long, Stephen H.; Say, Thomas; Adams, John (1823). Account of an expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and '20. 1. London: Longman, Hurst, Pees, Orre & Brown. pp. 168–174. 
  15. ^ Mussulman, Joseph (November 2004). "Thomas Say, Canis latrans". Discovering Lewis & Clark. Retrieved January 15, 2013. 
  16. ^ a b Clavijero, Francisco Javier; Cullen, Charles (1817). The history of Mexico : collected from Spanish and Mexican historians, from manuscripts and ancient paintings of the Indians : together with the conquest of Mexico by the Spaniards : illustrated by engravings with critical dissertations on the land, the animals, and inhabitants of Mexico. 1. Philadelphia: Thomas Dobson. p. 57. OCLC 13601464. 
  17. ^ Bullock, W. (1824). Six months' residence and travels in Mexico: containing remarks on the present state of New Spain, its natural productions, state of society, manufactures, trade, agriculture, and antiquities, &c. : with plates and maps. London: John Murray, Albemarle-Street. pp. 119, 261. 
  18. ^ a b c d e f Seton 1909, p. 789
  19. ^ a b c d e f g h i j k l Gier, H.T. (1974). "Ecology and Behavior of the Coyote (Canis latrans)". In Fox, M. W. The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution. New York: Van Nostrand Reinhold. pp. 247–262. ISBN 978-0-442-22430-1. OCLC 1093505. 
  20. ^ a b c d e f Bekoff, M.; Gese, E. M. (2003). "Coyote (Canis latrans)". In Feldhamer, G. A.; Thompson, B. C.; Chapman, J. A. Wild Mammals of North America: Biology, Management, and Conservation (2 ed.). Baltimore, Maryland: Johns Hopkins University Press. pp. 467–481. ISBN 978-0-8018-7416-1. OCLC 51969059. 
  21. ^ a b c d e f g h i j k Young & Jackson 1978, pp. 6–7
  22. ^ Curtis, E. S. (1928). The North American Indian. Volume 18 – The Chipewyan. The Western woods Cree. The Sarsi. Classic Books Company. p. 201. ISBN 978-0-7426-9818-5
  23. ^ a b Crawford, J. M. (1989). Cocopa Dictionary. University of California Press. p. 445. ISBN 978-0-520-09749-0. OCLC 20012309. 
  24. ^ a b LeClire, N.; Cardinal, G. (1998). Alberta Elders' Cree Dictionary. University of Alberta. p. 279. ISBN 978-0-88864-284-4. OCLC 659111819. 
  25. ^ a b Martin, J. P.; Mauldin, M. M. (2004). A Dictionary of Creek/Muskogee. University of Nebraska Press. p. 153. ISBN 978-0-8032-8302-2. OCLC 43561668. 
  26. ^ a b Albert, R.; Shaul, D. L. (1985). A Concise Hopi and English Lexicon. John Benjamins Publishing. p. 26. ISBN 978-90-272-2015-8. OCLC 777549431. 
  27. ^ Bright, William; Gehr, Susan. "Dictionary entry for coyote". Karuk Dictionary and Texts. Karuk Tribe & UC Berkeley. Retrieved May 22, 2015. 
  28. ^ a b c Reid, F. A. (2009). A Field Guide to the Mammals of Central America and Southeast Mexico. Oxford University Press. p. 259. ISBN 978-0-19-534322-9. OCLC 237402526. 
  29. ^ Aoki, Haruo (1994). Nez Percé dictionary. University of California Press. p. 491. ISBN 978-0-520-09763-6. OCLC 463788185. 
  30. ^ Neundorf, A. (1983). A Navajo/English Bilingual Dictionary: Áłchíní Bi Naaltsoostsoh. University of New Mexico Press. p. 512. ISBN 978-0-8263-3825-9. OCLC 57357517. 
  31. ^ Quintero, C. (2004). Osage Grammar. University of Nebraska Press. p. 83. ISBN 978-0-8032-3803-9. OCLC 57614396. 
  32. ^ Parks, R. P.; Pratt, L. N. (2008). A Dictionary of Skiri Pawnee. University of Nebraska Press. p. 119. ISBN 978-0-8032-1926-7. OCLC 940905155. 
  33. ^ a b c Dayley, J. P. (1989). Tümpisa (Panamint) Shoshone Dictionary. University of California Press. p. 436. ISBN 978-0-520-09754-4. OCLC 489876664. 
  34. ^ a b Pitkin, H. (1985). Wintu Dictionary. University of California Press. pp. 65, 573. ISBN 978-0-520-09613-4. OCLC 12313411. 
  35. ^ "Dictionary entry for coyote". Yurok Language Project. UC Berkeley. Retrieved May 22, 2015. 
  36. ^ Brewer, Douglas J.; Terence, Sir Clark; Phillips, Adrian (March 2002). Dogs in Antiquity: Anubis to Cerberus: The Origins of the Domestic Dog. Aris & Phillips. ISBN 978-0-85668-704-4. OCLC 49351443. 
  37. ^ a b Nowak, R. M. (1978). "Evolution and taxonomy of coyotes and related Canis". In Bekoff, M. Coyotes: Biology, Behavior, and Management. New York: Academic Press. pp. 3–16. ISBN 978-1-930665-42-2. OCLC 52626838. 
  38. ^ a b Lindblad-Toh, K.; Wade, C. M.; Mikkelsen, T. S.; Karlsson, E. K.; Jaffe, D. B.; Kamal, M.; Clamp, M.; Chang, J. L.; Kulbokas, E. J.; Zody, M. C.; Mauceli, E.; Xie, X.; Breen, M.; Wayne, R. K.; Ostrander, E. A.; Ponting, C. P.; Galibert, F.; Smith, D. R.; Dejong, P. J.; Kirkness, E.; Alvarez, P.; Biagi, T.; Brockman, W.; Butler, J.; Chin, C. W.; Cook, A.; Cuff, J.; Daly, M. J.; Decaprio, D.; et al. (2005). "Genome sequence, comparative analysis and haplotype structure of the domestic dog". Nature. 438 (7069): 803–819. Bibcode:2005Natur.438..803L. doi:10.1038/nature04338. ISSN 1476-4687. OCLC 889377779. PMID 16341006. 
  39. ^ a b von Holdt, Bridgett M.; Cahill, James A.; Fan, Zhenxin; Gronau, Ilan; Robinson, Jacqueline; Pollinger, John P.; Shapiro, Beth; Wall, Jeff; Wayne, Robert 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. doi:10.1126/sciadv.1501714. ISSN 2375-2548. OCLC 892343396. 
  40. ^ Kurtén, Björn (1980). Pleistocene Mammals of North America. Columbia University Press. pp. 167–9. ISBN 978-0-231-03733-4. OCLC 5830693. 
  41. ^ Tedforda, Richard H.; Wang, Xiaoming; Taylor, Beryl E. (2009). "Phylogenetic Systematics of the North American Fossil Caninae (Carnivora: Canidae)". Bulletin of the American Museum of Natural History (325): 1–218. doi:10.1206/574.1z (inactive 2016-08-22). ISSN 1937-3546. OCLC 46381488. 
  42. ^ a b c Meachen, J. A.; Samuels, J. X. (2012). "Evolution in coyotes (Canis latrans) in response to the megafaunal extinctions". Proceedings of the National Academy of Sciences. 109 (11): 4191. doi:10.1073/pnas.1113788109. ISSN 1091-6490. OCLC 475396714. PMID 22371581. 
  43. ^ a b Meachen, J. A.; Janowicz, A. C.; Avery, J. E.; Sadleir, R. W. (2014). "Ecological Changes in Coyotes (Canis latrans) in Response to the Ice Age Megafaunal Extinctions". PLoS ONE. 9 (12): e116041. Bibcode:2014PLoSO...9k6041M. doi:10.1371/journal.pone.0116041. PMID 25551387. 
  44. ^ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 532–628. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  45. ^ Young & Jackson 1978, p. 249
  46. ^ a b c d e f g h i Merriam, C. H. (1897). "Revision of the coyotes or prairie wolves, with descriptions of new forms". Proceedings of the Biological Society of Washington. 11: 19–33. 
  47. ^ a b c d e f g h i j k l m n o p q r Nowak, Ronald M. (1979). "History and Statistical Analysis of Recent Populations". In Wiley, E. O. North American Quaternary Canis. Lawrence, Kansas: University of Kansas Printing Service. pp. 9–10. 
  48. ^ a b Elliot, D. G. (1904). The land and sea mammals of Middle America and the West Indies, vol. II, pp. 467–8. Chicago.
  49. ^ a b Nelson, E. W. (1932). "Remarks on coyotes, with description of a new subspecies from Salvador". Proceedings of the Biological Society of Washington. 45: 223–225. 
  50. ^ a b Méndez-Carvajal, P. (2014). "Mammalia, Carnivora, Canidae, Canis latrans (Say, 1823): actual distribution in Panama". Check List. 10 (2): 376–379. doi:10.15560/10.2.376. ISSN 1809-127X. OCLC 828104819. 
  51. ^ Merriam, C. H. (1904). "A new coyote from southern Mexico". Proceedings of the Biological Society of Washington. 17: 157. Archived from the original on September 7, 2014. 
  52. ^ Goldman, E. A. (1936). "A new coyote from Honduras". Jour. Washington Acad. Sci. 26: 32–34. 
  53. ^ Young & Jackson 1978, p. 263
  54. ^ a b Townsend, C. H. (1912). "Mammals collected by the 'Albatross' expedition in Lower California in 1911, with descriptions of new species". Bulletin of the American Museum of Natural History. 31: 117–130. 
  55. ^ Bailey, V. (1905). "Biological survey of Texas". North American Fauna. 25: 1–222. doi:10.3996/nafa.25.0001. ISSN 1944-4575. OCLC 273060038. 
  56. ^ a b Jackson, H. H. T. (1949). "Two new coyotes from the United States". Proceedings of the Biological Society of Washington. 62: 31–32. 
  57. ^ a b c d Young & Jackson 1978, pp. 121–24
  58. ^ Cartaino 2011, pp. 61–3
  59. ^ Fox 1978, p. 105
  60. ^ Fox 1978, p. 136
  61. ^ Anderson, T. M.; Vonholdt, B. M.; Candille, S. I.; Musiani, M.; Greco, C.; Stahler, D. R.; Smith, D. W.; Padhukasahasram, B.; Randi, E.; Leonard, J. A.; Bustamante, C. D.; Ostrander, E. A.; Tang, H.; Wayne, R. K.; Barsh, G. S. (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. ISSN 1095-9203. OCLC 34298537. PMC 2903542free to read. PMID 19197024. 
  62. ^ Zimmer, Carl (January 21, 2013). "Snow Coyotes and Spirit Bears". National Geographic Magazine. Retrieved July 1, 2016. 
  63. ^ a b c Way, J.G.; Rutledge, L.; Wheeldon, T.; B.N. White (2010). "Genetic characterization of Eastern "Coyotes" in eastern Massachusetts" (PDF). Northeastern Naturalist. 17 (2): 189–204. doi:10.1656/045.017.0202. ISSN 1938-5307. JSTOR 40664873. OCLC 46381506. 
  64. ^ 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. ISSN 1938-5307. JSTOR 4499900. OCLC 46381506. 
  65. ^ a b Bekoff, M. (1978). "Behavioral Development in Coyotes and Eastern Coyotes". Coyotes: Biology, Behavior, and Management. New York: Academic Press. pp. 97–127. ISBN 978-1-930665-42-2. OCLC 52626838. 
  66. ^ Hilton, Henry (1978). "Systematics and Ecology of the Eastern Coyote". In Bekoff, M. Coyotes: Biology, Behavior, and Management. New York: Academic Press. pp. 210–28. ISBN 978-1-930665-42-2. OCLC 52626838. 
  67. ^ Vonholdt, B. M.; Pollinger, J. P.; Earl, D. A.; Knowles, J. C.; Boyko, A. R.; Parker, H.; Geffen, E.; Pilot, M.; Jedrzejewski, W.; Jedrzejewska, B.; Sidorovich, V.; Greco, C.; Randi, E.; Musiani, M.; Kays, R.; Bustamante, C. D.; Ostrander, E. A.; Novembre, J.; Wayne, R. K. (2011). "A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids". Genome Research. 21 (8): 1294–1305. doi:10.1101/gr.116301.110. ISSN 1549-5469. OCLC 37589079. PMC 3149496free to read. PMID 21566151. 
  68. ^ a b Fox, M. W. (1974). "Evolution of Social Behavior in Canids". The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution. New York: Van Nostrand Reinhold. pp. 429–60. ISBN 978-0-442-22430-1. OCLC 1093505. 
  69. ^ Bekoff, Marc; Diamond, Judy (1976). "Precopulatory and copulatory behavior in coyotes". Journal of Mammalogy. 57 (2): 372–375. doi:10.2307/1379696. ISSN 0022-2372. JSTOR 1379696. OCLC 1800234. 
  70. ^ Gese, Eric M., and Robert L. Ruff. "Scent-marking by coyotes, Canis latrans: the influence of social and ecological factors." Animal Behaviour 54.5 (1997): 1155-1166.
  71. ^ Mech, D. L. (2003). The Wolves of Minnesota: Howl in the Heartland. Voyageur Press. p. 75. ISBN 978-0-89658-509-6. OCLC 43694482. 
  72. ^ Hennessy, C. A.; Dubach, J.; Gehrt, S. D. (2012). "Long-term pair bonding and genetic evidence for monogamy among urban coyotes (Canis latrans)". Journal of Mammalogy. 93 (3): 732. doi:10.1644/11-MAMM-A-184.1. ISSN 1545-1542. OCLC 39098574. 
  73. ^ Fox 1978, p. 33
  74. ^ Mlot, Chris (1998). "The Coyotes of Lamar Valley: In Yellowstone, the master adapter learns to deal with wolves". Science News. 153 (5): 76–78. doi:10.2307/4010114. JSTOR 4010114. 
  75. ^ Young & Jackson 1978, pp. 82–90
  76. ^ Asa, C. S.; Mech, D. (1995). "A review of the sensory organs in wolves and their importance to life history". In Carbyn, L. D.; Fritts, S. H.; Seip, D. R. Ecology and Conservation of Wolves in a Changing World. Edmonton, Alberta: University of Alberta. pp. 287–291. ISBN 978-0-919058-92-7. OCLC 35162905. 
  77. ^ Österholm, H. (1964). "The significance of distance reception in the feeding behaviour of fox (Vulpes vulpes L.)". Acta Zoologica Fennica. 106: 1–31. 
  78. ^ Wells, M. C. (1978). "Coyote senses in predation – environmental influences on their relative use". Behavioural Processes 3. 3 (2): 149–158. doi:10.1016/0376-6357(78)90041-4. PMID 24924653. 
  79. ^ Wells, M. C.; Lehner, P. N. (1978). "Relative importance of distance senses in coyote predatory behavior". Animal Behaviour. 26: 251–258. doi:10.1016/0003-3472(78)90025-8. 
  80. ^ Young & Jackson 1978, pp. 91–92
  81. ^ Young & Jackson 1978, p. 97
  82. ^ Young & Jackson 1978, p. 98
  83. ^ a b c Young & Jackson 1978, pp. 93–96
  84. ^ Saunders, D.A. "Adirondack Ecological Center: Coyote". College of Environmental Science and Forestry, SUNY. 
  85. ^ Brundige, G. C. (1993). "Predation ecology of the eastern coyote Canis latrans "var.", in the central Adirondacks, New York". State University of New York, College of Environmental Science and Forestry, Syracuse. 
  86. ^ Boyer, R. H. (1949). "Mountain coyotes kill yearling black bear in Sequoia National Park". Journal of Mammalogy. 30: 75. doi:10.1093/jmammal/30.1.75. ISSN 1545-1542. OCLC 39098574. 
  87. ^ Way, J. G.; Horton, J. (2004). "Coyote kills harp seal" (PDF). Canid News. 7 (1). ISSN 1545-1542. OCLC 39098574. Archived from the original (PDF) on May 13, 2006. 
  88. ^ Klauber, Lawrence Monroe (1997). Rattlesnakes: Their Habits, Life Histories, and Influence on Mankind. 1 (2 ed.). Berkeley, California: University of California Press. pp. 1072–4. ISBN 978-0-520-21056-1. OCLC 39523012. 
  89. ^ Cordey, Huw (2013). North America: A World in One Continent. Philadelphia: Running Press. ISBN 978-0-7624-4843-2. OCLC 808413615. 
  90. ^ Berger, K. M.; Gese, E. M. (2007). "Does interference competition with wolves limit the distribution and abundance of coyotes?". Journal of Animal Ecology. 76 (6): 1075–1085. doi:10.1111/j.1365-2656.2007.01287.x. PMID 17922704. 
  91. ^ Stains, H. J. (1974). "Distribution and Taxonomy of the Canidae". In Fox, M. W. The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution. New York: Van Nostrand Reinhold. pp. 3–26. ISBN 978-0-442-22430-1. OCLC 1093505. 
  92. ^ Sargeant, Alan B.; Allen, Stephen H. (1989). "Observed interactions between coyotes and red foxes". Journal of Mammalogy. 70 (3): 631–633. doi:10.2307/1381437. ISSN 1545-1542. JSTOR 1381437. OCLC 39098574. 
  93. ^ a b Fedriani, J. M.; Fuller, T. K.; Sauvajot, R. M.; York, E. C. (2000). "Competition and intraguild predation among three sympatric carnivores" (PDF). Oecologia. 125 (2): 258–270. doi:10.1007/s004420000448. ISSN 1432-1939. OCLC 76327396. PMID 24595837. Archived from the original (PDF) on October 6, 2011. 
  94. ^ Hornocker, M.; Negri, S. (2009). Cougar: Ecology and Conservation. University of Chicago Press. p. 170. ISBN 978-0-226-35347-0. OCLC 609634655. 
  95. ^ Litvaitis, J. A.; D. J. Harrison (1989). "Bobcat-coyote niche relationships during a period of coyote population increase". Canadian Journal of Zoology. 67 (5): 1180–1188. doi:10.1139/z89-170. 
  96. ^ a b Bunnell, Kevin D.; Flinders, Jerran T.; Wolfe, Michael L. (2006). "Potential Impacts of Coyotes and Snowmobiles on Lynx Conservation in the Intermountain West". Wildlife Society Bulletin. 34 (3): 828–838. doi:10.2193/0091-7648(2006)34[828:PIOCAS]2.0.CO;2. ISSN 1938-5463. JSTOR 3784713. OCLC 60353682. 
  97. ^ a b Gipson, P. S.; Kamler, J. F (2002). "Bobcat Killed by a Coyote". The Southwestern Naturalist. 47 (3): 511–513. doi:10.2307/3672519. ISSN 0038-4909. JSTOR 3672519. OCLC 525604174. 
  98. ^ Knick, S. T. (1990). "Ecology of bobcats relative to exploitation and a prey decline in southeastern Idaho". Wildlife Monographs. 108 (108): 1–42. ISSN 1938-5455. JSTOR 3830671. OCLC 60618095. 
  99. ^ a b Ripple, W. J.; Wirsing, A. J.; Beschta, R. L.; Buskirk, S. W. (2011). "Can restoring wolves aid in lynx recovery?" (PDF). Wildlife Society Bulletin. 35 (4): 514. doi:10.1002/wsb.59. 
  100. ^ O’Donoghue, M.; Hofer, E. J.; Doyle, F. I. (1995). "Predator versus predator". Natural History. 104: 6–9. 
  101. ^ Rockwood, Larry L. (2015). Introduction to Population Ecology. Chichester, United Kingdom: John Wiley and Sons. p. 273. ISBN 978-1-118-94755-5. OCLC 899267730. 
  102. ^ Bushkirk, S. W.; Ruggiero, L. F.; Krebs, C. J. (2000). "Habitat Fragmentation and Interspecific Competition: Implications for Lynx Conservation". In Ruggiero, L. F.; Aubry, K. B.; Buskirk, S. W.; Koehler, G. M.; Krebs, C. J.; McKelvey, K. S.; Squires, J. R. Ecology and conservation of lynx in the United States (PDF). Denver: University of Colorado Press. pp. 91–92. 
  103. ^ Hidalgo-Mihart, M. G. (2004). "Historical and present distribution of coyote (Canis latrans) in Mexico and Central America". Journal of Biogeography. 31 (12): 2025–2038. doi:10.1111/j.1365-2699.2004.01163.x. 
  104. ^ De la Rosa, C. L.; Nocke, C. C. (2010). "Carnivore Evolution: Central America and the Great North-South Migrations". A Guide to the Carnivores of Central America: Natural History, Ecology, and Conservation. University of Texas Press. ISBN 978-0-292-78951-7. 
  105. ^ a b Silver, H.; Silver, W. T. (1969). "Growth and Behavior of the Coyote-like Canid of Northern New England and Observations on Canid Hybrids". The Wildlife Society, Wildlife Monographs. 17: 24–25. ISSN 1938-5455. JSTOR 3830473. OCLC 60618095. 
  106. ^ Fox 1978, pp. 134–135
  107. ^ Bee, James (1981). Mammals in Kansas. University of Kansas. p. 165. 
  108. ^ Michael A. Mares; Oklahoma Museum of Natural History (Norman, Okla.) (1999). Encyclopedia of Deserts. University of Oklahoma Press. pp. 137–8. ISBN 978-0-8061-3146-7. 
  109. ^ Lehner, Philip N. (1978). "Coyote Communication". In Bekoff, M. Coyotes: Biology, Behavior, and Management. New York: Academic Press. pp. 127–162. ISBN 978-1-930665-42-2. OCLC 52626838. 
  110. ^ Young & Jackson 1978, pp. 107–114
  111. ^ a b Gier, H. T.; Kruckenberg, S. M.; Marler, R. J. (1978). "Parasites and diseases of coyotes". In Bekoff, M. Coyotes: biology, behavior, and management. New York: Academic Press. pp. 37–71. ISBN 978-1-930665-42-2. OCLC 52626838. 
  112. ^ Chai, J. Y.; Darwin, Murrell K.; Lymbery, A. J. (2005). "Fish-borne parasitic zoonoses: Status and issues". International Journal for Parasitology. 35 (11–12): 1233–1254. doi:10.1016/j.ijpara.2005.07.013. PMID 16143336. 
  113. ^ a b Watts, L. S. (2006). Encyclopedia of American Folklore. Infobase Publishing. pp. 93–94. ISBN 978-1-4381-2979-2. OCLC 465438817. 
  114. ^ a b Schwartz, M. (1998). A History of Dogs in the Early Americas. Yale University Press. pp. 146–149. ISBN 978-0-300-07519-9.
  115. ^ a b Kelley, D. H. (1955). "Quetzalcoatl and his Coyote Origins". El México Antiguo. 8: 397–416. 
  116. ^ Gillespie, Angus K.; Mechling, Jay (1987). American Wildlife in Symbol and Story. University of Tennessee Press. pp. 225–230. ISBN 978-0-87049-522-9. OCLC 14165533. 
  117. ^ Way, J. G. (2012). "Love wolves and hate coyotes? A conundrum for canid enthusiasts" (PDF). International Wolf. 22 (4): 8–11. Archived from the original (PDF) on December 24, 2012. 
  118. ^ Harris, M. (1979). Cultural Materialism: The Struggle for a Science of Culture. New York: AltaMira Press. pp. 200–1. ISBN 978-0-7591-0135-7. OCLC 47100657. 
  119. ^ Lynch, P. A.; Roberts, J. (2010). Native American Mythology A to Z. Infobase Publishing. p. 27. ISBN 978-1-4381-3311-9. OCLC 720592939. 
  120. ^ Miller, M. E.; Taube, K. A. (1993). The Gods and Symbols of Ancient Mexico and the Maya: An Illustrated Dictionary of Mesoamerican Religion. Thames and Hudson. p. 92. ISBN 978-0-500-05068-2. OCLC 27667317. 
  121. ^ Olivier, G. (2003). Mockeries and Metamorphoses of an Aztec God: Tezcatlipoca, "Lord of the Smoking Mirror". University Press of Colorado. p. 32. ISBN 978-0-87081-745-8. OCLC 52334747. 
  122. ^ Dell'Amore, Christine (March 2006). "City Slinkers". Smithsonian. Retrieved June 14, 2012. 
  123. ^ a b c d "Coyote Attacks: An Increasing Suburban Problem" (PDF). March 2004. Archived from the original (PDF) on September 26, 2007. Retrieved August 19, 2007. 
  124. ^ a b Baker, Rex O. (2007). "A Review of Successful Urban Coyote Management Programs Implemented to Prevent or Reduce Attacks on Humans and Pets in Southern California". Wildlife Damage Management Conferences – Proceedings: 382–392. 
  125. ^ Kays, R.; Curtis, A.; Kirchman, J. J. (2009). "Rapid adaptive evolution of northeastern coyotes via hybridization with wolves" (PDF). Biology Letters. 6 (1): 89–93. doi:10.1098/rsbl.2009.0575. PMC 2817252free to read. PMID 19776058. 
  126. ^ a b c "Coyote Predation – Description". A. Wade, Dale & E. Bowns, James. Procedures for Evaluating Predation on Livestock and Wildlife. Archived from the original on August 6, 2007. Retrieved August 19, 2007. 
  127. ^ "Sheep and Goats Death Loss" (PDF). National Agricultural Statistics Service. May 6, 2005. Retrieved December 27, 2007. 
  128. ^ "Sheep and Lamb Predator and Nonpredator Death Loss in the United States, 2015" (PDF). United States Department of Agriculture. 2015. Retrieved July 1, 2016. 
  129. ^ "Sheep and lamb inventory". United States Department of Agriculture. Retrieved February 1, 2010. 
  130. ^ "Controlling wily coyotes? Still no easy answers". NBC News. December 7, 2009. Retrieved September 14, 2013. 
  131. ^ Collinge, Mark; Timm, R. M.; Madon, M. B. (2008). "Relative risks of predation on livestock posed by individual wolves, black bears, mountain lions and coyotes in Idaho". Proceedings of the Vertebrate Pest Conference: 129–133. Archived from the original on February 23, 2015. 
  132. ^ "Livestock Protection Dogs" (PDF). Wildlife Services. October 2010. Retrieved July 3, 2016. 
  133. ^ "Livestock guarding dogs fact sheet". Animal and Plant Health Inspection Service United States Department of Agriculture. Retrieved April 3, 2012. 
  134. ^ "Ranchers' Guide to Wolf Depredation". Montana State University. 2006. Archived from the original on April 9, 2013. Retrieved July 1, 2016. 
  135. ^ Hardesty, Greg (May 5, 2005). "For coyotes, pets are prey". Greg Hardesty. Orange County Register. Archived from the original on July 15, 2007. 
  136. ^ "A coyote attacks in Weymouth and kills a dog". WHDH-TV – New England News. May 14, 2007
  137. ^ Macur, Juliet (2010). "Coyote vs. Greyhound: The Battle Lines Are Drawn". New York Times. Retrieved July 3, 2016. 
  138. ^ a b Young & Jackson 1978, pp. 115–116
  139. ^ Seton 1909, p. 816
  140. ^ Young & Jackson 1978, pp. 119–21
  141. ^ a b Young & Jackson 1978, pp. 64–9
  142. ^ Schultz, J. W. (1962). Blackfeet and Buffalo: Memories of Life Among the Indians. University of Oklahoma Press. pp. 141–3. ISBN 978-0-8061-1700-3. OCLC 248716. 
  143. ^ Etter, J. (February 15, 1998). "Coyote Blends In as Best Bird Dog for Durham Man". The Oklahoman. Retrieved July 1, 2016. 

Bibliography[edit]

Further reading[edit]

Books
Video
  • Shelly, Priya (June 2016). Living with Coyote (18 minutes). Aeon (digital magazine). "Wildlife management is really a misnomer. It’s about managing people." Short documentary aiming to address the problem of coyotes preying on rural livestock and scavenging in urban and suburban areas. The video combines interviews (with ranchers, coyote scientists/ researchers, etc), animation and scenery from the western US, aiming to discuss North America’s coyote dilemma, and to present several different views and perspectives on the debate on how human activities and attitudes may impact the problem.

External links[edit]