Coyote

• 1. Introduction

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  In 2026, a coyote, Canis latrans, swam to the southern edge of Alcatraz Island. To get there, it crossed more than a mile of water from the nearest land in San Francisco. A swim like that would surprise anyone who pictures the coyote only as a lean shape on a dry western ridge.

  Surprise is the coyote's specialty. This North American canine is smaller than the gray wolf, yet it adapts to landscapes reshaped by people. Urban coyotes are common in many cities.

  How did a modest predator become so adaptable? Why do geneticists find coyote DNA in most North American wolves? And why do the stories people tell about this animal swing between reverence and contempt?

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• 2. Names and First Descriptions

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  The word coyote began as coyōtl in Nahuatl, passed into Spanish, and eventually reached English. Its first appearance in print came in 1780, in the Historia de México by the historian Francisco Javier Clavijero. English readers met it in 1824, in William Bullock's Six months' residence and travels in Mexico. Bullock spelled it cayjotte and cocyotie, and only by the 1880s did the spelling settle as coyote. Pronunciation never settled. Speakers in eastern states, near the Mexican border, and outside the United States tend to say three syllables, while those in western and central states say two.

  The scientific name, Canis latrans, means barking dog, a nod to the many sounds the animal makes. Everyday English offers prairie wolf, brush wolf, cased wolf, little wolf, and American jackal. Native languages offer far more. The Navajo say Ma'ii, the Chinook say Italipas, and the Pawnee say Ckirihki.

  On the 5th of May 1805, in northeastern Montana, Meriwether Lewis described an animal he called the "small wolf or burrowing dog of the prairies". He noted that it moved in bands of ten or twelve and dug burrows near places where game passed often. He also heard it bark exactly like a small dog, and he described its eyes as deep sea green, small, and piercing. European traders on the upper Missouri already knew the animal well.

  Thomas Say gave the coyote its formal scientific description in September 1819. He was at the site of Lewis and Clark's Council Bluffs, 15 miles up the Missouri River from the mouth of the Platte. A government-sponsored expedition under Major Stephen Long had brought him there. Say carried the first edition of the Lewis and Clark journals, which included Biddle's edited version of Lewis's coyote notes. His account appeared in 1823. On the next page of his journal he described a second animal, which he named Canis nubilus, the Great Plains wolf. Say was the first person to record the difference between the two.

  Earlier writers had not been so careful. A priest in Kaskaskia, Illinois, wrote in 1750 of local "wolves" that were smaller and less daring than European ones. Historians cannot always tell which animal such early writers meant.

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• 3. Lighter Than Wolves

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  A male coyote usually weighs 8 to 20 kg, and a female 7 to 18 kg, though size depends on where the animal lives. Northern subspecies average 18 kg, while the coyotes of southern Mexico average 11.5 kg. Total length runs from 1.0 to 1.35 m on average, including a tail of about 40 cm. The largest coyote on record was a male killed near Afton, Wyoming, on the 19th of November 1937. It measured 1.5 m from nose to tail and weighed 34 kg.

  Coyote fur is mostly light gray and red or fulvous, mixed with black and white. Coyotes at high elevations tend toward black and gray, while desert coyotes look more fulvous or whitish-gray. The coat has a short, soft underlayer and long, coarse guard hairs. Northern animals grow longer and denser fur, and some Mexican and Central American forms have fur that is almost bristly.

  Between March 1938 and June 1945, federal and cooperative hunters killed 750,000 coyotes. Only two of them showed traits consistent with albinism, which makes the condition extremely rare.

  Beside a gray wolf, a coyote shows longer ears, a thinner frame, face, and muzzle, and a relatively larger braincase. Its coat varies much less than a wolf's. Its tail hangs downward when it walks or runs, where a wolf holds its tail horizontal. Its tracks are more elongated and less rounded than a dog's.

  The family tree stretches back millions of years. Xiaoming Wang and Richard Tedford, leading authorities on carnivore evolution, proposed that the genus Canis descended from a coyote-like animal called Eucyon davisi. Canis remains first appear about 6 million years ago, in the Miocene, in the southwestern United States and Mexico. By about 5 million years ago the larger Canis lepophagus lived there, and by about 1 million years ago the coyote itself existed. Wang and Tedford saw this as a straight line of descent.

  The modern coyote still carries marks of that heritage. Its skull and jaws are narrower than a wolf's, and its sagittal crest is low or flat, which points to a weaker bite. Its molars have larger chewing surfaces, reflecting a diet that leans on plants. In these respects it resembles the fox-like ancestors of the genus more than the wolf does.

  The oldest fossils within the modern coyote's range come from Hamilton Cave in West Virginia, dated to 0.74-0.85 million years ago. Pleistocene coyotes, known as C. l. orcutti, were larger and sturdier than today's, with teeth built for shearing meat. They shrank within 1,000 years of the Quaternary extinction event, when their large prey died out. The dire wolf vanished too, but gray wolves quickly took over its big-game niche and likely killed off the large coyotes. Natural selection then favored the slender modern form.

  In 1993, a study noted that the skulls of North American wolves resemble the coyote more closely than the skulls of Eurasian wolves do. DNA would later explain why.

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• 4. Wolf Blood

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  In 2016, a whole-genome study proposed that all North American wolves and coyotes split from a common ancestor about 51,000 years ago. That date clashes with a coyote-like specimen found in strata about 1 million years old. The same study found wolf ancestry in every coyote it examined, and coyote ancestry in every North American wolf. It also found that the red wolf and eastern wolf are mixtures of gray wolf and coyote in different proportions.

  A 2018 study compared specimens from across the whole range and produced the numbers. A polar wolf from Greenland and a coyote from Mexico were the purest animals in the sample. Coyotes from Alaska, California, Alabama, and Quebec carried almost no wolf ancestry. Those from Missouri, Illinois, and Florida showed 5-10% wolf ancestry. Red wolves were 40% wolf and 60% coyote. Eastern timber wolves were 60% wolf and 40% coyote, and Great Lakes wolves were 75% wolf and 25% coyote. Mexican wolves and Atlantic Coast wolves carried 10% coyote ancestry, Pacific Coast and Yellowstone wolves 5%, and Canadian archipelago wolves less than 3%.

  Another 2018 whole-genome comparison turned up a stranger result. The common ancestor of coyotes and gray wolves apparently mixed with a "ghost" population, an extinct and unidentified canid. This ghost was genetically close to the dhole and arose after the African wild dog had split from the other canids. Researchers proposed that the coyote sits at the base of the tree because it kept more of the ghost canid's mitochondrial genome.

  The eastern coyote of northeastern North America shows mixing in action. It probably arose after gray and eastern wolves were wiped out in the northeast, letting coyotes move into former wolf ranges and mate with the survivors. Female eastern coyotes weigh 21% more than male western coyotes. Eastern pups have longer legs by the age of 35 days, and their teeth come in later and in a different order. They reach sexual maturity at two years, much later than western coyotes. They also fight less, and their pups fight only after play begins, the reverse of the western pattern.

  The red wolf may owe about 75-80% of its genome to the coyote, after its declining numbers in the Southeastern Woodlands forced wolf-coyote mating and repeated backcrossing with local coyotes. The eastern wolf probably came from a wolf-coyote mixture that was then extensively backcrossed with gray wolves.

  Coyotes also mate with domestic dogs, producing animals called coydogs, but this is rare in the wild. The mating cycles of dogs and coyotes do not coincide, and coyotes are usually hostile to dogs. It happens mainly where coyotes are expanding into areas with few of their own kind. Pup survival is lower, because dogs do not form pair bonds with coyotes. Hybrids are fertile and can be bred through four generations.

  Dogs have left visible marks on coyote coats. Black melanistic coyotes owe their pelts to a mutation that first arose in domestic dogs. A population of white, non-albino coyotes in Newfoundland owes its color to a melanocortin 1 receptor mutation inherited from Golden Retrievers.

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• 5. Pups and Pairs

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  Coyote families form in midwinter, when females enter estrus. Pair bonding can begin two to three months before mating, and the mating tie lasts 5 to 45 minutes. A female in heat advertises with scent marking and increasingly frequent howling. A single female can draw up to seven reproductive males, which may trail her for as long as a month. Once she chooses, the rejected males do not interfere and move on. Unlike wolves, coyotes are strictly monogamous, even where food is plentiful and coyotes are dense.

  The new pair claims a territory and either digs a den or cleans out an abandoned badger, marmot, or skunk burrow. Dens have been found under abandoned homestead shacks, grain bins, drainage pipes, and railroad tracks. During pregnancy the male often hunts alone and carries food back to the female. She may line the den with dried grass or with fur pulled from her own belly. Gestation lasts 63 days, and the average litter is six, though food supply and local density change that number. A den can have several entrances and passages, and one den may serve for years. If fleas invade, the female moves the pups.

  Newborn pups weigh 200 to 500 g and depend entirely on milk for their first 10 days. Their eyes open at about 10 days, and the incisors come in at about 12. Parents start offering regurgitated solid food after 12-15 days. Canines erupt at 16 days, the second premolars at 21, and walking begins at about 20 days. Running comes at six weeks.

  Play among coyote pups begins with fighting, unlike wolf pups, which play first. By three weeks, coyote pups bite each other with less restraint than wolf pups do. By four to five weeks they have settled their dominance order and play more than they fight. One favorite move is the "hip-slam". The father helps feed, groom, and guard the litter, but he abandons it if the mother disappears before weaning is finished.

  By June to July the family leaves the den, and the pups follow their parents on patrol and on hunts. Some pups leave in August, though others stay much longer. They reach adult dimensions at eight months and adult weight a month after that. Not every coyote lives in a family. Bachelor males, nonreproductive females, and young adults may form temporary packs to keep company or to take down prey too large for one animal. Females that fail to mate sometimes help their sisters or mothers raise pups.

  Feeding territories vary enormously, from 0.4 to 62 square kilometres. Coyotes mark them by urinating with a raised leg and scratching the ground. Outside the denning season they rarely defend them, and intruders are usually chased and sparred with rather than killed.

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• 6. Hunters and Howlers

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  Despite the popular belief that smell guides the hunt, two experiments on red foxes and coyotes found that visual cues matter most. A coyote needs an estimated 600 g of food a day, or 250 kg a year, not counting insects, fruit, and grass. About 90% of its diet is meat. Mouse-sized rodents are caught by pouncing, and ground squirrels are chased down. Small prey is usually taken by a single animal.

  Large prey calls for teamwork. A breeding pair does most of the work, pursuing a large ungulate, hamstringing it, and harassing it until it falls. Snow depth and crust density strongly affect success. Younger animals usually stay out of these hunts, and coyotes cache surplus food like other canids.

  Coyotes have been seen killing porcupines in pairs, flipping them onto their backs with their paws and attacking the soft belly. Only old, experienced coyotes manage it, and young ones who try often end up injured by quills. Rattlesnakes are killed too, partly for food and partly to protect pups at the den. The coyote teases the snake until it stretches out, then bites its head and shakes it.

  A jar found in Mexico, dated to 1250-1300 CE, shows coyotes and American badgers working together. The partnership is real, since the two species sometimes help each other dig out rodents. Some coyotes have been seen resting their heads on badger companions or licking their faces without protest.

  The menu is remarkably broad. Coyotes eat bison, mostly as carrion, along with deer, elk, moose, rabbits, birds, lizards, snakes, fish, and insects. They also eat strawberries, blackberries, peaches, apples, watermelons, and carrots, and in winter and early spring they eat large amounts of grass. In Death Valley, they gorge on hawkmoth caterpillars or beetles during the spring flowering months. Coyotes in coastal California now eat more marine food than their ancestors did, which is thought to follow the disappearance of the grizzly bear there. They are also picky, since shrews, moles, and brown rats appear in their diet far less than their numbers would predict.

  One description calls the coyote "the most vocal of all wild North American mammals". Adults make at least 11 distinct sounds, grouped as agonistic and alarm calls, greetings, and contact calls. Woofs are low-intensity threats or alarms near the den, and they send pups scurrying into their burrows. Huffs are sharp, high-intensity threats made by rapidly expelling air. Yelps signal submission.

  Greetings are gentler. Submissive animals give low-frequency whines, often with tail wagging and muzzle nibbling. The "wow-oo-wow" has been called a greeting song. When pack members reunite, they may erupt in a group yip howl, perhaps the last step of an elaborate greeting ceremony. The lone howl, the coyote's most iconic sound, may announce an individual that has been separated from its pack.

  Adult fights are quieter than all that. Combatants approach with tails waving and jaws open, snarling, and the fights are typically silent. Males tend to rise into a vertical stance, while females fight on all four paws.

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• 7. Spreading Across a Continent

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  In the 1940s, wolves colonized Isle Royale, and the island's coyotes were driven to extinction. The coyote's range across the continent shifted the same way when wolves were removed. Before European colonization its range was limited to the Southwest, the Plains, and northern and central Mexico. During the 19th century it moved north and east, and it spread further after 1900 as land was converted and wolves were exterminated. In Yellowstone National Park, the coyote population in the Lamar River Valley fell by 39% after wolves were reintroduced in the 1990s. In Grand Teton National Park, coyotes are 33% fewer where wolves live.

  A 2017 genetic study suggests that coyotes were not originally found in the eastern United States at all. From the 1890s, dense forests became farmland and wolf control was widespread, leaving a vacant niche. Two genetically distinct populations moved in. The first dispersal began in the early 20th century from the northern Great Plains. It reached New England through the northern Great Lakes and southern Canada, and Pennsylvania through the southern Great Lakes. The two streams met in New York and Pennsylvania in the 1940s.

  The second dispersal began in the mid-20th century in Texas and reached the Carolinas in the 1980s. Both streams grew rapidly, and they are forecast to meet along the mid-Atlantic coast. The study suggests that long-distance dispersal, gene flow from local populations, and fast population growth may be linked.

  The push south continued through the late 1970s, south of Costa Rica, and the early 1980s, in northern Panama. It followed cattle pastures spreading into tropical rain forest. Many believed coyotes were recent arrivals in southern Mexico and Central America, but Pleistocene and Early Holocene records show they were there long before. In January 2013, a coyote was recorded in the Chepo District of eastern Panama, beyond the Panama Canal. Some fear a spread into South America if the Pan-American Highway ever closes the Darién Gap.

  Competitors push back. In the eastern Sierra Nevada, coyotes compete with cougars over mule deer, and cougars usually dominate and sometimes kill them. Wolves tolerate no coyotes nearby, though coyotes sometimes trail wolves to feed on their kills. When a wolf or an adult cougar shows up at a carcass, the coyote usually flees.

  Bobcats and lynx fare worse against coyotes. The biologist Stanley Paul Young never once saved a trapped bobcat from being killed by coyotes, and he described two incidents in which coyotes chased bobcats up trees. In central Alberta, lynx are more abundant where coyotes are few. Because the coyote is so widespread and abundant, the International Union for Conservation of Nature lists it as least concern.

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• 8. Pest and Trickster

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  In 2004, the National Agricultural Statistics Service attributed 60.5% of 224,000 sheep deaths from predation to coyotes. Because coyotes are far more numerous and widespread than wolves, they cause more overall losses. United States government agents shoot, poison, trap, and kill about 90,000 coyotes a year to protect livestock. In Utah, more than 11,000 coyotes were killed for bounties totaling over $500,000 in the fiscal year ending the 30th of June 2017. Guard dogs have worked well, and a 1986 survey found that 82% of sheep producers who used them called the dogs an economic asset.

  There have been only two confirmed fatal attacks on people. The victims were three-year-old Kelly Keen in Glendale, California, and the nineteen-year-old singer-songwriter Taylor Mitchell in Nova Scotia, Canada. In the 30 years leading up to March 2006, at least 160 attacks occurred in the United States, mostly in Los Angeles County. Verified attacks numbered 41 in 1988-1997 and 48 in 1998-2003. Urban coyotes are losing their fear of people, especially where people feed them. Some have chased joggers and bicyclists, and a few have targeted small children.

  About three to five pets attacked by coyotes arrive each week at the Animal Urgent Care hospital of South Orange County, California. Most are dogs, since cats rarely survive. At one Southern California site, coyotes ate most of a colony of feral cats and then kept returning for the cat food that people left out.

  For the fur trade, the coyote was worthless until the mid-19th century. By 1860, with beavers in decline, wolfers on the Great Plains earned 75 cents to $1.50 per coyote skin. In the early 1950s pelts sold for $5 to $25. Most went into trimmings such as coat collars, and some were dyed black to imitate silver fox.

  In the Maidu creation story, Coyote introduces work, suffering, and death to the world. Zuni lore has him bringing winter by stealing light from the kachinas. In a Tohono O'odham flood story, he helps Montezuma survive a flood that destroys humanity. The Navajo Coyote brings death, explaining that without it there would be no room to plant corn. In the Crow creation story, Old Man Coyote is the Creator himself. Across many nations of the Southwest and Plains, Coyote appears as a trickster who rebels against convention with deception and humor. After colonization, Anglo-American tales recast him as cowardly and untrustworthy, and unlike the gray wolf, his public image has not recovered.

  Before the Spanish conquest of the Aztec Empire, the coyote carried real weight in Mesoamerica. Warriors at Classic-era Teotihuacan dressed in coyote costumes to draw on its predatory power. Aztec mythology had Huehuecóyotl, meaning "old coyote", the god of dance, music, and carnality, drawn in several codices as a man with a coyote's head. In some tales he seduced Xochiquetzal, the goddess of love, and so brought war into the world. The epigrapher David H. Kelley argued that Quetzalcoatl grew out of older Uto-Aztecan images of the coyote as mankind's "Elder Brother", a creator and culture hero linked to the morning star.

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