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==Behavior== ===Social and reproductive behaviors=== [[File:Gpa bill coyote pups 3.jpg|thumb|Mearns' coyote (''C. l. mearnsi'') pups playing]] [[File:Pair of Coyotes Playing in Santa Teresa County Park (45917548654).jpg|thumb|The "hip-slam"<ref name="fox136"/> is a common play behavior]] [[File:Pack of coyotes on snow.jpg|thumb|A pack of coyotes in [[Yellowstone National Park]]]] Like the Eurasian golden jackal, the coyote is gregarious, but not as dependent on [[Biological specificity|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.<ref name="fox1974">{{cite book|last=Fox|first=M. W. |year=1974|chapter=Evolution of Social Behavior in Canids|pages=429β60|title=The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution|location=New York|publisher=Van Nostrand Reinhold|isbn=978-0-442-22430-1|oclc=1093505}}</ref> 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 on their own. 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]].<ref name="gier1974"/> Pair bonding can occur 2β3 months before actual [[Copulation (zoology)|copulation]] takes place.<ref>{{cite journal|last1=Bekoff|first1=Marc|first2=Judy|last2=Diamond |year=1976| title=Precopulatory and copulatory behavior in coyotes|journal=Journal of Mammalogy|volume=57|issue=2|pages=372β375|doi=10.2307/1379696|issn=0022-2372|oclc=1800234|jstor=1379696}}</ref> The [[copulatory tie]] can last 5β45 minutes.<ref>{{cite journal | last1 = Carlson | first1 = Debra A. | last2 = Gese | first2 = Eric M. | year = 2008 | title = Reproductive biology of the coyote (Canis latrans): integration of mating behavior, reproductive hormones, and vaginal cytology | journal = Journal of Mammalogy | volume = 89 | issue = 3| pages = 654β664 | doi=10.1644/06-mamm-a-436r1.1| pmid = 32287378 | pmc = 7108653 | doi-access = free }}</ref> A female entering estrus attracts males by scent marking<ref name="Gese1997">{{cite journal | last1 = Gese | first1 = Eric M. | last2 = Ruff | first2 = Robert L. | year = 1997 | title = Scent-marking by coyotes, Canis latrans: the influence of social and ecological factors | journal = Animal Behaviour | volume = 54 | issue = 5| pages = 1155β1166 | doi=10.1006/anbe.1997.0561| pmid = 9398369 | citeseerx = 10.1.1.540.1024 | s2cid = 33603362 }}</ref> and howling with increasing frequency.<ref name=bekoff2003/> 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.<ref name="gier1974"/> Unlike the wolf, which has been known to practice both [[Monogamy in animals|monogamous]] and bigamous matings,<ref>{{cite book|last=Mech| first=D. L. |year=2003|title=The Wolves of Minnesota: Howl in the Heartland|publisher=Voyageur Press|page=75|isbn=978-0-89658-509-6|oclc=43694482}}</ref> the coyote is strictly monogamous, even in areas with high coyote densities and abundant food.<ref>{{Cite journal | doi = 10.1644/11-MAMM-A-184.1|url=http://www.mammalsociety.org/articles/long-term-pair-bonding-and-genetic-evidence-monogamy-among-urban-coyotes-canis-latrans| title = Long-term pair bonding and genetic evidence for monogamy among urban coyotes (''Canis latrans'')| journal = Journal of Mammalogy| volume = 93| issue = 3| pages = 732β742| year = 2012| last1 = Hennessy | first1 = C. A. | last2 = Dubach | first2 = J. | last3 = Gehrt | first3 = S. D. |issn=1545-1542|oclc=39098574| doi-access = free}}</ref> 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 [[American badger|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.<ref name="gier1974"/> 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.<ref name="bekoff2003"/> Coyote pups are born in dens, hollow trees, or under ledges, and weigh {{convert|200|to|500|g|lb|abbr=on}} at birth. They are [[altricial]], and are completely dependent on [[milk]] for their first 10 days. The [[incisor]]s erupt at about 12 days, the [[canine teeth|canine]]s at 16, and the second [[premolar]]s 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 12β15 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.<ref name="gier1974"/> Unlike wolf pups, coyote pups begin seriously fighting (as opposed to play fighting) prior to engaging in play behavior. A common play behavior includes the coyote "hip-slam".<ref name="fox136"/> 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.<ref name="fox33">{{Harvnb|Fox|1978|p=33}}</ref> The male plays an active role in feeding, [[social grooming|grooming]], and guarding the pups, but abandons them if the female goes missing before the pups are completely [[Weaning|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.<ref name="gier1974"/> ===Territorial and sheltering behaviors=== {{image frame |caption=[[Scent marking]] |content= {{CSS image crop |Image = LA_River_Camera_Project_(30730459477).jpg |bSize = 450 |cWidth = 210 |cHeight = 170 |oTop = 70 |oLeft = 130 }}{{CSS image crop |Image = Coyote_pair_in_Wheeling_%288530485951%29.jpg |bSize = 500 |cWidth = 210 |cHeight = 130 |oTop = 170 |oLeft = 200 }}}} Individual feeding territories vary in size from {{convert|0.4|to|62|km2|sqmi|abbr=on|sigfig=2}}, 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,<ref name="gier1974"/> and is much less aggressive towards intruders than the wolf is, typically chasing and sparring with them, but rarely killing them.<ref>{{cite journal|last=Mlot|first=Chris|year=1998|title=The Coyotes of Lamar Valley: In Yellowstone, the master adapter learns to deal with wolves|journal=Science News|volume=153 |issue=5|pages=76β78|doi=10.2307/4010114|jstor=4010114}}</ref> Conflicts between coyotes can arise during times of food shortage.<ref name="gier1974"/> Coyotes mark their territories by [[raised-leg urination]] and ground-scratching.<ref name="WellsBekoff1981"/><ref name="Gese1997"/> 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 [[canyon]]s, [[Washout (erosion)|washout]]s, [[coulee]]s, [[Bank (geography)|bank]]s, 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.<ref name="young82">{{Harvnb|Young|Jackson|1978|pp=82β90}}</ref> A single den can be used year after year.<ref name="bekoff2003"/> ===Hunting and feeding behaviors=== While the popular consensus is that [[olfaction]] is very important for hunting,<ref>{{cite book|last1=Asa|first1=C. S.|last2=Mech|first2=D.|year=1995|chapter=A review of the sensory organs in wolves and their importance to life history |title=Ecology and Conservation of Wolves in a Changing World|editor1-last=Carbyn|editor1-first=L. D.|editor2-last=Fritts|editor2-first=S. H.|editor3-last=Seip|editor3-first=D. R.|location=Edmonton, Alberta|publisher= University of Alberta|pages=287β291|isbn=978-0-919058-92-7|oclc=35162905}}</ref> 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<ref>{{cite journal|last=Γsterholm|first=H.|year=1964|title=The significance of distance reception in the feeding behaviour of fox (''Vulpes vulpes L.'')|journal=Acta Zoologica Fennica|volume=106|pages=1β31}}</ref> and coyotes.<ref>{{cite journal|last=Wells|first=M. C.|year=1978|title=Coyote senses in predation β environmental influences on their relative use|journal=Behavioural Processes|volume=3|issue=2|pages=149β158 |doi=10.1016/0376-6357(78)90041-4|pmid=24924653|s2cid=22692213}}</ref><ref>{{cite journal|last1=Wells|first1=M. C.|last2=Lehner|first2=P. N.|year=1978|title=Relative importance of distance senses in coyote predatory behavior |journal= Animal Behaviour|volume=26|pages=251β258|doi=10.1016/0003-3472(78)90025-8|s2cid=53204333}}</ref> {{image frame|border=no|content={{photo montage|size=220 |photo1a=Coyote Pouncing.jpg |photo1b=Leaping Coyote Seedskadee NWR (16117597568).jpg |photo2a=Coyote Hunting Rodents in Santa Teresa County Park (30035278974).jpg |photo2b=Coyote (Canis latrans) (7147080735).jpg |border=0 |color=transparent |text=Coyotes pouncing on prey }} {{photo montage|size=220 |photo1a=Coyote - Dead Elk (4634125254).jpg |photo1b=Coyote eating bison YNP.jpg |border=0 |color=transparent |text=Coyotes with elk and bison carcasses }}}} When hunting large prey, the coyote often works in pairs or small groups.<ref name="bekoff1977"/> Success in killing large [[ungulate]]s 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.<ref name="bekoff2003"/> The coyote pursues large prey, typically [[hamstringing]] the animal, and subsequently then harassing it until the prey falls. Like other canids, the coyote [[Hoarding (animal behavior)|cache]]s excess food.<ref name="young91">{{Harvnb|Young|Jackson|1978|pp=91β92}}</ref> Coyotes catch mouse-sized rodents by pouncing, whereas [[ground squirrel]]s are chased. Although coyotes can live in large groups, small prey is typically caught singly.<ref name="bekoff2003"/> Coyotes have been observed to kill [[North American porcupine|porcupine]]s 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.<ref name="young97">{{Harvnb|Young|Jackson|1978|p=97}}</ref> Coyotes sometimes [[urination|urinate]] on their food, possibly to claim ownership over it.<ref name="WellsBekoff1981">Wells, Michael C., and Marc Bekoff. "[http://animalstudiesrepository.org/cgi/viewcontent.cgi?article=1035&context=acwp_ena An observational study of scent-marking in coyotes, Canis latrans]." (1981).</ref><ref name="young98">{{Harvnb|Young|Jackson|1978|p=98}}</ref> Recent evidence demonstrates that at least some coyotes have become more nocturnal in hunting, presumably to avoid humans.<ref>{{Cite news|url=https://www.nytimes.com/2018/06/15/science/animals-human-nocturnal-study.html |archive-url=https://ghostarchive.org/archive/20220101/https://www.nytimes.com/2018/06/15/science/animals-human-nocturnal-study.html |archive-date=2022-01-01 |url-access=limited |title=Mammals Go Nocturnal in Bid to Avoid Humans|newspaper=The New York Times |date=June 15, 2018 |access-date=2018-06-21|last1=Jacobs |first1=Julia }}{{cbignore}}</ref><ref>{{Cite journal |last=Gaynor |date=2021-06-15 |title=The influence of human disturbance on wildlife nocturnality |journal=Science |volume=360 |issue=6394 |pages=1232β1235 |doi=10.1126/science.aar7121 |pmid=29903973 |s2cid=49212187 |doi-access=free }}</ref> Coyotes may occasionally form [[mutualism (biology)|mutualistic]] hunting relationships with [[American badger]]s, assisting each other in digging up rodent prey.<ref>{{cite web|author=U.S. Fish and Wildlife Service|title=Spotted! A Coyote and Badger Hunting Together|year=2016|url=https://www.fws.gov/news/blog/index.cfm/2016/11/2/Spotted-A-Coyote-and-Badger}}</ref> 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 jar found in Mexico dated to 1250β1300 [[Current era|CE]] depicting the relationship between the two.<ref name="young93"/> Food scraps, pet food, and animal feces may attract a coyote to a trash can.<ref>{{Cite web|title=Why is there a coyote in my yard? Food lures and other answers|url=https://www.humanesociety.org/resources/why-there-coyote-my-yard-food-lures-and-other-answers|website=The Humane Society of the United States|access-date=2020-05-07}}</ref> ===Communication=== [[File:Howl (cropped).jpg|thumb|A coyote [[howling]]]] [[File:Pack of coyotes howling.ogg|thumb|Pack of coyotes howling at night]] ====Body language==== Being both a gregarious and solitary animal, the variability of the coyote's visual and vocal repertoire is intermediate between that of the solitary foxes and the highly social wolf.<ref name="fox1974"/> 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.<ref name="silver1969">{{cite journal|author1=Silver, H. |author2= Silver, W. T. |jstor=3830473|title=Growth and Behavior of the Coyote-like Canid of Northern New England and Observations on Canid Hybrids|year= 1969|journal=The Wildlife Society, Wildlife Monographs|volume= 17|issue= 17 |pages= 24β25| issn=1938-5455 |oclc=60618095}}</ref> 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.<ref name="fox134">{{Harvnb|Fox|1978|pp=134β135}}</ref> 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.<ref name="silver1969"/> ====Vocalizations==== [[File:Yelping Coyote.webm|thumb|A yelping coyote]] The coyote has been described as "the most vocal of all [wild] North American mammals".<ref name="Mammals in Kansas">{{cite book|last1=Bee|first1=James|title=Mammals in Kansas|date=1981|publisher=University of Kansas|page=165}}</ref><ref name="Mares_Oklahoma_1999">{{cite book|author1=Michael A. Mares|author2=Oklahoma Museum of Natural History (Norman, Okla.)|title=Encyclopedia of Deserts|url=https://books.google.com/books?id=g3CbqZtaF4oC&pg=PA137|year=1999|publisher=University of Oklahoma Press|isbn=978-0-8061-3146-7|pages=137β8}}</ref> 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.<ref name="lehner1978"/> Growls are used as threats at short distances but have also been heard among pups playing and copulating males. Huffs are high-intensity threat vocalizations produced by rapid expiration of air. Barks can be classed as both long-distance threat vocalizations and alarm calls. Bark howls may serve similar functions. Yelps are emitted as a sign of submission, while high-frequency whines are produced by dominant animals acknowledging the submission of subordinates. Greeting vocalizations include low-frequency whines, 'wow-oo-wows', and group yip howls. Low-frequency whines are emitted by submissive animals and are usually accompanied by tail wagging and muzzle nibbling.<ref name="lehner1978"/> 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.<ref name="lehner1978">{{cite book|last=Lehner|first=Philip N. |year=1978|chapter=Coyote Communication|pages=127β162|editor-first=M. |editor-last=Bekoff|title=Coyotes: Biology, Behavior, and Management|publisher=Academic Press| location=New York|isbn=978-1-930665-42-2|oclc=52626838}}</ref>
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