The coyote (pronounced /kaɪˈoʊtiː, ˈkaɪ.oʊt/) (Canis latrans), the American jackal or the prairie wolf, is a species of canid found throughout North and Central America, ranging from Panama in the south, north through Mexico, the United States and Canada. It occurs as far north as Alaska and all but the northernmost portions of Canada. There are currently 19 recognized subspecies, with 16 in Canada, Mexico and the United States, and 3 in Central America. Unlike its cousin the Gray Wolf, which is Eurasian in origin, the coyote evolved in North America during the Pleistocene epoch 1.810 million years ago alongside the Dire Wolf. Unlike the wolf, the coyote's range has expanded in the wake of human civilization, and coyotes readily reproduce in metropolitan areas. It is thought by certain experts that the coyote's North American origin may account for its greater adaptability than the wolf, due to North America's greater prehistoric predation pressures.

Name

The name "coyote" is borrowed from Mexican Spanish, ultimately derived from the Nahuatl word cóyotl. Its scientific name, Canis latrans, means "barking dog."

Description

Coyote profile

The color of the coyote's pelt varies from grayish brown to yellowish gray on the upper parts, while the throat and belly tend to have a buff or white color. The forelegs, sides of the head, muzzle and paws are reddish brown. The back has tawn-colored underfur and long, black-tipped guard hairs that form a black dorsal stripe and a dark cross on the shoulder area. The black-tipped tail has a scent gland located on its dorsal base. Coyotes shed once a year, beginning in May with light hair loss, ending in July after heavy shedding. The ears are proportionately large in relation to the head, while the feet are relatively small in relation to the rest of the body. Certain experts have noted that the shape of a domestic dog's brain case is closer to the coyote's in shape than the wolf's. Mountain dwelling coyotes tend to be dark furred while desert coyotes tend to be more yellowish in color.

Coyotes typically grow up to 75–87 centimeters (30–34 inches) in length, not counting a tail of 30-40 cm (12-16 in), stand about 58-66 cm (23-26 in) at the shoulder and, on average, weigh from 7–21 kilograms (15–46 pounds). Northern coyotes are typically larger than southern subspecies, with the largest coyotes on record weighing 74¾ pounds (33.7 kg) and measuring over five feet in total length.

The coyote's dental formula is I 3/3, C 1/1, Pm 4/4, M usually 2/2, occasionally 3/3, 3/2, or 2/3 X 2 = 40, 42, or 44. Normal spacing between the upper canine teeth is 1⅛ to 1⅜ inches (29 to 35 mm) and 1 to 1¼ inches (25 to 32 mm) between the lower canine teeth.

The upper frequency limit of hearing for coyotes is 80 kHz, compared to the 60 kHz of domestic dogs.Compared to wolves, and similarly to domestic dogs, coyotes have a higher density of sweat glands on their paw pads. This trait, however, is absent in the large New England coyotes, which are thought to have some wolf ancestry.

During pursuit, a coyote may reach speeds up to 43 mph (69 km/h), and can jump a distance of over 4 meters (13⅛ feet).

Behavior

	Coyote in Yellowstone National Park

Though coyotes have been observed to travel in large groups, they primarily hunt in pairs. Typical packs consist of six closely related adults, yearlings and young. Coyote packs are generally smaller than wolf packs and associations between individuals are less stable, thus making their social behaviour more in line with that of the dingo. It has been theorized that this is due to an earlier expression of aggression, and the fact that coyotes reach their full growth in their first year, unlike wolves who reach it in their second. Common names of coyote groups are a band, a pack, or a rout. Coyotes are primarily nocturnal, but can often be seen during daylight hours. Coyotes were once essentially diurnal, but have adapted to more nocturnal behavior with pressure from humans (McClennen et al., 2001).

Coyotes are capable of digging their own burrows, though they often appropriate the burrows of groundhogs or American badgers. Coyote territorial ranges can be as much as 19 kilometers in diameter around the den, and travel occurs along fixed trails.

In areas where wolves have been exterminated, coyotes usually flourish. For example, as New England became increasingly settled and the resident wolves were eliminated, the coyote population increased, filling the empty biological niche. Coyotes appear better able than wolves to live among people.

Coyotes have been known to live a maximum of 10 years in the wild and 18 years in captivity.They seem to be better than dogs at observational learning.

Reproduction

Newborn coyote pup		Seven coyote pups

Female coyotes are monoestrous, and remain in heat for 2–5 days between late January and late March, during which mating occurs. Once the female chooses a partner, the mated pair may remain temporarily monogamous for a number of years. Depending on geographic location, spermatogenesis in males takes around 54 days, and occurs between January and February. The gestation period lasts from 60 to 63 days. Litter size ranges from 1 to 19 pups; the average is 6. These large litters act as compensatory measures against the high juvenile mortality rate, with approximately 50-70% of pups not surviving to adulthood.The pups weigh approximately 250 grams at birth, and are initially blind and limp-eared. Coyote growth rate is faster than that of wolves, being similar in length to that of the dhole. The eyes open and ears become erect after 10 days. Around 21–28 days after birth, the young begin to emerge from the den, and by 35 days they are fully weaned. Both parents feed the weaned pups with regurgitated food. Male pups will disperse from their dens between months 6 and 9, while females usually remain with the parents and form the basis of the pack. The pups attain full growth between 9 and 12 months. Sexual maturity is reached by 12 months. Unlike wolves, mother coyotes will tolerate other lactating females in their pack.

Interspecific Hybridization

	A Coyote-German Shepherd hybrid

Coyotes will sometimes mate with domestic dogs, usually in areas like Texas and Oklahoma where the coyotes are plentiful and the breeding season is extended because of the warm weather. The resulting hybrids, called coydogs, maintain the coyote's predatory nature, along with the dog's lack of timidity toward humans, making them a more serious threat to livestock than pure-blooded animals. This cross-breeding has the added effect of confusing the breeding cycle. Coyotes usually breed only once a year, while coydogs will breed year-round, producing many more pups than a wild coyote. Differences in the ears and tail are generally what can be used to distinguish coydogs from domestic/feral dogs or pure coyotes. Breeding experiments in Germany with poodles, coyotes, and later on with the resulting dog-coyote hybrids showed that unlike wolfdogs, coydogs show a decrease in fertility, significant communication problems as well as an increase of genetic diseases after three generations of interbreeding.

Coyotes have also been known on occasion to mate with wolves, though this is less common than with dogs due to the wolf's hostility to the coyote. The offspring, known as a coywolf, is generally intermediate in size to both parents, being larger than a pure coyote, but smaller than a pure wolf. A study showed that of 100 coyotes collected in Maine, 22 had half or more wolf ancestry, and one was 89 percent wolf. A theory has been proposed that the large eastern coyotes in Canada are actually hybrids of the smaller western coyotes and wolves that met and mated decades ago as the coyotes moved toward New England from their earlier western ranges. The Red Wolf is thought by certain scientists to be in fact a wolf/coyote hybrid rather than a unique species. Strong evidence for hybridization was found through genetic testing, which showed that red wolves have only 5% of their alleles unique from either Gray Wolves or coyotes. Genetic distance calculations have indicated that red wolves are intermediate between coyotes and Gray Wolves, and that they bear great similarity to wolf/coyote hybrids in southern Quebec and Minnesota. Analyses of mitochondrial DNA showed that existing Red Wolf populations are predominantly coyote in origin.

Communication

Hearing a coyote is much more common than seeing one. The calls a coyote makes are high-pitched and variously described as howls, yips, yelps and barks. These calls may be a long rising and falling note (a howl) or a series of short notes (yips). These calls are most often heard at dusk or night, but may be heard in the day. Although these calls are made throughout the year, they are most common during the spring mating season and in the fall when the pups leave their families to establish new territories.

Ecology

Diet and hunting

	Coyote on elk carcass in winter in Lamar Valley

Coyotes are opportunistic, versatile carnivores with a 90% mammalian diet, depending on the season. They primarily eat small mammals, such as voles, prairie dogs, eastern cottontails, ground squirrels, and mice, though they will eat birds, snakes, lizards, deer, javelina, and livestock, as well as large insects and other large invertebrates. Any species of birds that nests on the ground are targeted by coyotes. Though they will consume large amounts of carrion, they tend to prefer fresh meat. Part of the coyote's success as a species is its dietary adaptability. As such, coyotes have been known to eat human rubbish and domestic pets. Fruits and vegetables are a significant part of the coyote's diet in the autumn and winter months.

Coyotes shift their hunting techniques in accordance with their prey. When hunting small animals such as mice, they slowly stalk through the grass, and use their acute sense of smell to track down the prey. When the prey is located, the coyotes stiffen and pounce on the prey in a cat-like manner. Coyotes will commonly work in teams when hunting large ungulates such as deer. Coyotes may take turns in baiting and pursuing the deer to exhaustion, or they may drive it towards a hidden member of the pack. When attacking large prey, coyotes attack from the rear and the flanks of their prey. Occasionally they also grab the neck and head, pulling the animal down to the ground. Coyotes are persistent hunters, with successful attacks sometimes lasting as much as 21 hours; even unsuccessful ones can continue more than 8 hours before the coyotes give up. Depth of snow can affect the likelihood of a successful kill. Packs of coyotes can bring down prey as large as adult elk, which usually weigh over 250 kg (550 lbs).

The average distance covered in a night's hunting is 4 km (2½ mi).

Interspecific predatory relationships

		Rolf Peterson investigating a coyote carcass killed by a wolf in Yellowstone National Park, January 1996

The gray wolf is a significant predator of coyotes wherever their ranges overlap. Since the Yellowstone Gray Wolf Reintroduction in 1995 and 1996, the local coyote population went through a dramatic restructuring. Until the wolves returned, Yellowstone National Park had one of the densest and most stable coyote populations in America due to a lack of human impacts. Two years after the wolf reintroductions, the pre-wolf population of coyotes had been reduced 50% through both competitive exclusion and intraguild predation. In Grand Teton, coyote densities were 33% lower than normal in the areas where they coexisted with wolves, and 39% lower in the areas of Yellowstone where wolves were reintroduced. In one study, about 16% of radio-collared coyotes were preyed upon by wolves. Yellowstone coyotes have had to shift their territories as a result, moving from open meadows to steep terrain. Carcasses in the open no longer attract coyotes; when a coyote is chased on flat terrain, it is often killed. They feel more secure on steep terrain where they will often lead a pursuing wolf downhill. As the wolf comes after it, the coyote will turn around and run uphill. Wolves, being heavier, cannot stop and the coyote gains a large lead. Though physical confrontations between the two species are usually dominated by the larger wolves, coyotes have been known to attack wolves if they outnumber them. Both species will kill each other's pups given the opportunity. Wolf urine has been marketed and claimed to be an organic coyote deterrent, such as for deterring attacks on sheep.

Cougars sometimes kill coyotes. The coyote's instinctive fear of cougars has led to the development of anti-coyote sound systems which repel coyotes from public places by replicating the sounds of a cougar. Bears can also rarely kill coyotes, more likely in competitive rather than predatory attacks. However, both cougars and bears have been displaced from carcasses by coyote packs.

In sympatric populations of coyotes and red foxes, fox territories tend to be located largely outside of coyote territories. The principal cause of this separation is believed to be active avoidance of coyotes by the foxes. Interactions between the two species vary in nature, ranging from active antagonism to indifference. The majority of aggressive encounters are initiated by coyotes, and there are few reports of red foxes acting aggressively toward coyotes except when attacked or when their pups were approached. Conversely, foxes and coyotes have sometimes been seen feeding together.

Coyotes will sometimes form a symbiotic relationship with American badgers. Because coyotes are not very effective at digging rodents out of their burrows, they will chase the animals while they are above ground. Badgers on the other hand are not fast runners, but are well-adapted to digging. When hunting together, they effectively leave little escape for prey in the area.

In some areas, coyotes share their ranges with bobcats. It is rare for these two similarly sized species to physically confront one another, though bobcat populations tend to diminish in areas with high coyote densities. Coyotes (both single individuals and groups) have been known to occasionally kill bobcats, but in all known cases, the victims were relatively small specimens, such as adult females and juveniles.

Coyotes have also competed with and occasionally eaten Canadian lynxes in areas where both species overlap.