Mammatian Species Canis latrans. By Mare Bekoff . 79, pp. 1-9, 6 figs. Published 15 June 1977 by The American Society of Mammalogists Canis latrans Say, 1823 Coyote Canis tatrans Say, in James, 1823-168. Type locality Engineer Cantonment, about 19,2 km SE present town of Bir. Washington Co, Nebraska Canis ociropus Exchschoke 1829"). Type locality Sacramento River Valley near Sacramento, California, Lyeiscus cagotts Hamilton Sith (1839-168). Type locality Rio Frio. west slope of Mount Iztaccthuatl Mex. Gani rrr Woon 1950-187, Type city Re fork of the Arkansas iver, probably near 97 west longitude, near Perkins, Payne Co. tnaw Cimarron River, Oklahoma Canis testes Merriam (169135). Tope localty Tayabe Mountains ‘near Cloverdale, Nye Co., Nevada. Canis mearnsi Mentiam (163429). Type locality Quitobaquito, Pima Co, Arizona rey aa Canis microdon Merriam (1997:29. Type locality Mie conkers Tope ea ‘ Canis peninslae Merriam (1897:28) Type locality Senta Anita, ‘Cape Saint Leas, Baja Califor Canis estor Mernam (18973). Type locality Noland’s Rench, ‘San Juan River Valley, San Juan Co., Utah ani iis Mera (18730. Type lacaity ear Mansa, Canis lepticus Blot (1903:225), Type locality San Pedro Mart Mountain, 2,0 m, Baja California. Canis impavias Allen (19032600). Type lveality Rio dees Bocas, 32138 m, northwestern Durango. (Cans goldinant Mei (904187. Type leas San Vicente, Cant james Townsend (1912190. Type lvaty Tiburon aad, Canis hondurensis Goldman (1936:33), Type locality Cerro ‘Guinote, NE of Archaga, on the ‘elangs road north of ‘Teguetgalpa, Honduras. CONTEXT AND CONTENT. Order Carnivora, Family Caniae, Genus Canis, ia which, thee re iit ecognined species, There are 19 recognized. subspecies af C. larans, wthich is considered to bora’ cloe relative af the Jackal (C; aureus, Comesomelas, and C. udusius)ace Karten, 1918, an ‘Remaris. For more deiail on subspecies see Nelo 193), Jacke fon (951) and Hall and Kelson (1099) . .laorans Say (in James, 1823:168), see above (pallidus Mer sam and nebracenats Merriam are synonyms) . Lockropus Eschecholty(1829:1), tee above. E:cagotis (HHamiton-Smith, 1839:168), see above, Gt frtray Woodhouse -S1-180, ce above ‘ on Rio 1 Testes Mersiam (1897225), see above, C. 1. mearnsi Merriam (189129), see above (stor Merviam a ‘synonym U'nevadon Merriam (1997.20), see above U peninculae Merriam (1897:28), see above 1, bail Merziam (1897-33), see above I elepricus Et (1908-225), see above 1. impaticus Allen (1905-603), see above. F goldmant Merriam (1904 18), see above. 1 texensis Bailey (1905:175). Type locality Santa Gertrudis, Kieberg.Co., Texas 1. Jament Townsend (1912:180), tee above, I iokeyi Nelson (1982:229), Type locality ‘3.2 km W'Rio Gosscoran, La Union (03% Salvador 1. colatus Hall (1984:269). Type locality Isaaes Lake, Bowron Ieake Region, British Colua C.. hondarensis Goldman (1986:38), see shove C1. thamnos Jackson (198931) Type locality Basswood Island, 1 eal lla Aaland Co Wie tbe C. 1. umpquenss Jackson (1989:31). Type locality 5 mi , Douglas Co., Oregon DIAGNOSIS. The coyote can be differentiated from other Canis in the Wester Hemiaphere (gray wolf. lupus; red wall, AD appanse 1 Cerra Mogote, north Taide), C, rafus; and domestic dog, C. fanilaris) using a number of teria The coyote is typically smaller than the aray wall Gee General Characters and sso Mech, 1973), but there iy overlap vthen comparing the covote withthe red wolf and the domestic tog. Also. depending on geographic locale, there may be light ovgap wih Tapas Lammene and Bonet OS The ne pad of the coyote lapproximately 25 mm in diametes is smal iat of the wen nthe date of the pao the hind foot less than 32 mm as opposed to greater than 3B mim. re spectively). The ears of the coyote ate longer than those of the tay wll, The track of the coyote (approximately 10 mm by 60 mm—O. J. Murie, 1984) is more elongated than that of the domestic dog, but shorter than that of both the grey wolf and the red wolf Riley and McBride (1975) presented mean values Of 66 mm (37 to 72mm) and 102 mam (9 to 127 mm) for the track Tength (fromthe back of the heel to the ‘end. of the longest claw) for the coyote and red wolf. respectively. The stride, of the coyote i fess than that of the gray wolf or red wall, The mean length of the stride of the coyote is approx ately 414 mm (S24 to 483 mmly whereas that of rufus is proximately 658 mm (52 to 762 mms Riley and MeBride, 1885. Dental characters also have been used to distinguish atrans, lupus, and rufus, but Jackson (195) stressed that such measure: iments may not be especialy reliable. In the covote, the Ups oF the upper canine ecth usully extend below line drawn Uouutt the enteror mental foramina of the mandible when the mandibie is ariculated and the jaws closed: Howard 1919) iegested any for diterentatng ltrans fv fois that ut 95 reliable, depending on subspecies, A-ratio of palatal width (between the inne ara of vot of the Lipper fet molar) to length of the upper molar tothrow {trom the anterior margin f the aveolis of the Ft premolar to the posterior margin ofthe last molar alveaus) in calculated. Tf the Soar. tines the pall thy the specien 8 yo: if the ratio lees than 2, the specimen te 8 "Various cranial measurements have been used to diferen- late species of Canis (Lawrence and Bossert, 1967, 1969, 1975 Paradico, 1968; Paradiso and Nowak, 1971: Wortmann, 1971: ison ef al, 1974). The coyote has a relatively lager braincase than does C. tapus (Mech, 1974). Paradiso and Nowak did ex: tensive analyses on skulls of farransy lupus, and rufes, and {demonstrated the usefulness af indices, They found no overlay then comparing the largest coyote to the smallest wolf (upus) in" zygomatic ‘breadth (greatest distance across "zygomatal, st leat of the shal (see igure 1) ote ratio the rato SF the width across the outer edges of the sivcoll of the fnteriorTobes ofthe upper carmassale tothe length of the upper molar toothrow as defined sh ant specific ogntion. The differences between the red wolf andthe covate ave far grester than those between recognized subspecies of Ietrans. The mort rebale featore separating larant fr raf, {s'feacr sizer there in‘alimost'o overlap greatest length of the skall (igure 1. Also, rufus haa heavier bone structrey a Felutively broader skull, and generally a more pronounces sapital rest Lasrence_ and. Bomert (1982) using multple charscter ‘analyses and linear discrimination techniques, found nine cranial land six dental measurements (ace Lawrence and Bosser, 1967, table T ‘and "appesdin Ay Tor pariclar) that could be weed Felisbly to diflerratate datrans from lupus and femiliars but fo single character was found without overlap between pair OF species. C-latrons differed more from lupus and famiiars than aps difered from familiar “Te coyote bre difers from that of C. lupus (Radineky, 1973) in thatthe waif has'a dimple in the middle of the corona astus, whereas the coyote dows not, Using gross cerebellar Trorphsiogy, Atkins and Dillon (1971) distingushed fatran from both fapas‘and rufus sce Remarks). The coyote differs from Clapns tnd figs erkgcly Crone aa Wiens, 1938 Ste alo Seat, 197 Behaviorally, the coyote can be diferentated from C, lupus and €:faniliars. The coyote shows higher levels of aggression ler tn life than does the wolf or Beagle (and probally moatubitus yee! our) MAMMALIAN SPECIES 79, —ome es) a) —Com. (e) i _—_tomtan 0) —— mmr 110) m7) er) (Otlehome? omer en (aa Se) (Louisiono) nin. (Texos) Comer LATRANS thamnos aia (4) texensis a (12) smecensi ome 8) lateans ome 2) test an incototus oe 019) ttttt tt tt dodo 180 190 200 20 220 230 240 250 260 270 280 290 Fiovme 1 reatest length of skull (rm) in male gray wolves (lupus), red wolves (rufus) and coyotes (atrans) similar tend) line on top at fois evident in females (lender lin snge; black bar = 2 standard errors of the mean: vertical ‘mean: black bar plus white portion on elther side indicate one standard deviation on either side of the meat). After Paradiso and Nowak, 1971, with permission other domestic dogs as well) and also performs species ton appears to have occurred into Florida (Cunningham and Dun. {ppical” behavior called the “Inguinal response” mhich consists oF rotating «hind-eg outwards oF lifting it of the ground In Fesponse to ight inguinel contact (sce figure 2, and Bekoff 19a, 1973, 194 GENERAL CHARACTERS, The size of the covte varies with subspecies and geogr isis Hal tnd Kelson, 1950), Adult males usually are heavier and larger 'an_adult females (approximately 8 to 20 kg as oppored to 7 to 18 kg, reapectively)- In Sagehen Creek Basin (northeastern California), Hawthorne (1971) found that males averaged 11-12 Kg (Bi? to 1249 eg) and females averaged. 9.76 kg (7-72 to 12.5 kp), whereas in Texas Daniel (19754) found that 71 males averaged 16.75 kg, (11.35 to 20-48 kg] and 70 females averaged 18,62 eg 10.90 to 17.35 ks). ier (1975) reported that northern sepecies are larger (approximately 18 kg) than more southern Eubspecies on the Mexican deserts (1.5 bg). Body length varies {rom 1.0 to'1-35'm and the tal = about 400 mm long: Femeles are shorter than ‘males in both length and height. ‘he largest ‘coyote on record was taken in Wyoming. Ie weighed 38.98 ke and measured 1.60 m from Up of nose to tip of tal Voung, 1951) Color and texture of the for vary geographically. The hair Js longer and coarser in nanher subspecies (see Form). The banded nature of the hair presents blended colors gray mixed with 's reddish tint, Those coyotes at higher aldtudes tend Coward more gray and black, whereas those in the desert are ‘are fulvous (Jackson, 1951) Black patches may be found on the front of the foreteet and near the base and Up of the tal {ier 1968). ‘The belly and throat age paler than the rest af the body. Melanistic coyotes are rere (Young, 1951; van Wormer, 1968; Gipson, 1976) DISTRIBUTION. Coyotes are, Nearactic canide orig nally inhabiting open country and grasslands (Young, 1951; Gie, 1923). Within historic timey they have occupied many diverse habitats, They now can be found between 10” nor latitade (Conta Ria) snd 10° north latitude (northern Alaska) and through: Out the continental Unite Sater and Canada i, 3) The ange Of the coyote rs-expanding, When considering the expanding mnge of. larrans iis important to know whether this reflects true movement of coyotes or whether some recently discovered Populations Gn some cases afew dens) are the result of ani ‘mals being transplanted into the region by man. ‘Transplants: 1510) and Georgia (Fsher, 1975, Paradiso (1968) considered the expansion of the coyote into Arkansas, Mississippi, and Louisiana to have been unaesisted by hur FOSSIL RECORD. Fossils resembling C. latrans were found in Pleistocene deposits in Cumberland Cave, Maryland (Gidley, 1913: Matthew, 1930). Differentiation of modern canids feccurred in the Pleistocene and in Recent times (eee les, 1900: Colbert, 1969: Todd, 1970). Ficune 2, A “speciestypical” action, the inguinal response performed by the coyote onthe left The le isnot being pushed tip but is ited due to ight stimulation im the inguinal regionMAMMALIAN SPECIES 79 iGURE 3. Current distribution of coyote (T, probable trans- plants by man-see text. FORM. Coarse guard hairs are about 50 t9 90 mm long {in the mane, 8) to. 10 mim) with imbricate scales that are Scuminate nthe proximal region, ctenate medially and fattened distally. "The fine underfur has ‘coronal scales and may be fone as 50mm (see Adorjan and Kolenosky, 1969; Ogle and f ‘molt between late Faris, 1973). There ustally js one. main molt spring and autumn, front foot has five toes a foo here in dew caw pona mth ther fortes, ipso ermal” commana} abneryed specimen wh 4 ‘developed dew claw on cach hindigot. The toes are nor le dnd the stance digiigrade. There are 42 teeth (i 3B Ap ai me ter det er ef al.. 1974, and Bekoff and Jamiccon,. 1975). The Sku atu male weighs Detneen 170 anf 210 p ands 180 to 205 "mm long [igures 4 and’) fom the tip of the pre zsilla to the posterior nim of the coronal erest(Gier, 1968). Males show greater development of the sagittal ridge than do the females, "The structure and evolution of the coyote’s brain have been described by Radinsky (1969, 1973), Atking and Dillon 11971}. ‘and Ellas and Sehwarts (1971 Mossman and Duke (1973) described the ovaries of various ‘eanide. The adrenals of canids were studied by Ogle (971) and Heinrich (1072) ‘Ogle (L971) reported that the eft adrenal Iheavier than the sight in both males and females and that the Adrenals of females tend to he heavier than those of males GUTS 0.01 as opposed to 1:08 0.08 gs respectively. FUNCTION. The coyote’ furis similar in insulative value to that of the gray wolf TOple and Farr, 1943). The teal temperature of lerrans is “10°C (Os consumption = 7.35 mm tmine Shield, 19421. The longer winter far coat conserves heat ‘considerably better than the shorter summer ere of at 0 thermal coc ‘olfactory cues during thei aetivities and deposit “marks” {coven pcslly undue scereons) on conepcamas objects possibly Tor terry demarcation (as yet an unsubstantiated a Simption). Onoga and Harger 11966) reported that the coyotes -Promosilary sSagital Crest ntecpavietal, Fused wih te" Suprececipital ‘Lombdoidel Ridge [FIoURE 4, Skull of coyote: upper left, dorsal views lower lef, ‘ventral view (from Lecheitner, 1969, wth permission ofthe pub lisher they observed urinated an average, of once per 4.0 km and ‘deserted every 10-4 km, Gipson and Seslander (1972) reported Urination and defecation occurring three times per 1.6 km and fone ime per 3.2to 4. kim, respectively. Scratching of the ground iso common component a he emia. prove possible that scent from interdigital glands orate voeal repertoire (Tembrock, 1963; eliner, 1973). "The region of maximal sen sultry stm i 100 He to 30 Kite witha top lim ‘Petercen et a, 1968), The retina iz duplex and has a Ponderance of ds, The sbcolote scotopic (rod) threshold ape proximately 14 fogt-candles. The adaptation curve shows dstinet keane breaks (Horn and Lehner, 1973). The electocardiograph st the coyote based on the ventricular setivation process isthe same as of other carnivores [Stabuniewica, 1970 REPRODUCTION AND ONTOGENY. Field date on suman doth that courthip may begin as long ae2 to 8 mnths before re attempts mt copuistion. The female fs monoestrous, showing cr year. This usualy occurs between and March and bath males and females show seasonal ad gle oot anny gn hilogy itanicn, "1968. Kennelly, 1972: Dunber, 1978Sagital Postaitl Process ‘et rotted Portal" O ceria! Nowlory SDE trsit Se ramen Bement, \ Twit ca i i SA be S aaa tes Waa Freire tanbdoial “Peo | Hel nies FidGe™ External : ‘AicMory —Fatertital Process of Jugal Condyloid Process (> Coronsid Process \\ Conine Wenta! Foramina Ficume 5. Skull and jaw of coyote in lateral view (from Lechleitner, 1969, with permission of publisher. Proestrous lasts between 2 and 3. months (Whiteman, 1949: Kennelly and Roberts, 1969: Bekoff and Diamond, 1976) estrus laste aboat 2 to'5 days (Kleiman, 1908; personal observations), go eration secure about 213 days hare the end of the female receptivity. The times of, and durations of, proestrus and estrout’ differ in various locales sand the relationship. i fot clear-cut (Hamlet, 1938), The same is true for the spermator tenie eyele of the male. Bebavioral changes that wscur during Fourtship parallel these reported by Golani and Mendeleeah OAD for the golden jackal (C- aureus). Courtship “rituals” are not so highly pei that sian cannot cc, Cyto ‘Can tuceessfully mate with domestic doge, gray wolves, red ‘wolves, and jackals. Copulation ends wih the typical “copulatory the,” during which the males penis is locked in the female's vagina (Dewshury, 1972; Grandage, 1972; Fox and Bekoff, 1975) ‘The tie can last 15 to 28 minutes (Kleiman, 1968: Bekoff and Diamond, 1976). Iti generally accepted thet the same individoas will mate rom year to year, but not necessarily for ie ‘The percentage of females that breed during given year may vary irom 43% to. 90% depending on loeal conditions (Gier, 1968; Knowlton, 1972; Gipson eal, 1975). Both yearling ‘males and females are capable of reproducing, yearling females "sual breeding later than older female (Gee. 196) and thus Contributing minimally to the population (Knowlton, 1972), Cipson er al. (1975) found that no’ yearling females bred. However, {Gier, 1968) reported that in good rodent years, 15% of yearling females may breed. Nelli and Keith (1976) reported pregnancy rates of S4% for adult females and 14% for yearlings, wth adults shoving more (6.0) placental sears than yearlings (S.2)—aot a Significant difference). Asdell (19), summarizing the iterate, Feported the mean number of embryos in 1370 cases to be 6.23 land the number of den young in 1882 eases to be 5.70. Ham- Jett (1938) reported that the number of young per iter was about 85% of the number of embryos per female. Gier (1915) tstimated that the number of young born was equal to about ‘BOE of the ovulated ova depending on whether er not it was ‘good rodent year. Gipson et al. (1973) eported the mean nunn- ber of ova per breeding female to be 6.2, with 4.3 becoming implanted. Gestation laste approximately 68 days (58 to 65). Coyotes and dogs appear to develop similarly un utero” (Gier, 1068), ‘Average litter sie is six with aveex ratio of females to tales of about Tcl. Nels and Keith (1970) reported a mean litter size of 5.8 for 26 liters. Litter size i known to be accepted by population density (Knowlton, 1972} and odent populations (Cie, 1908) Liter size averaged 4.3 at high denaities and 0.9 at low densities. In good todent years mean litter size was 5.8 10 6:2 and in poorer rodent years average liter size was 4.4 105. MAMMALIAN SPECIES 79 ‘TaBLe 1. Mortality in coyote populations. (After Mathwig, 1973: 184; mean percentage values based on seven studies), ‘Aue (years) 1 41 Percent of population Young are horn blind and helpless, usually in an excavated nn Favorable den sites include brush covered slopes, steep ks, thickets, hollow logs, rock Iedges, often on south-facing slopes (Ger, 1968) Dens of ther animals may be used, Dene are ‘sually about 0.3m in diameter and may be from 1.5 to 1 Tong (van Wormer, 1508) More than ne entrance may fi tilitate the movement of young when the den site i= disturbed. ‘The same den may be ured from year to year, and dens may be' shared. Nels "and Keith (1970) found that 3-of 29 dens had two litters, ‘Young are nursed by their mother (sce Ewer, 1978:331 for data onthe composition of eoyote milk) and are weaned at aout week 5 to 7 (Snow, 1907: unpublished deta). They begin to eat Solid food at about week 3, and the female (and possibly the ‘male) begins regurgitating semi-solid food at this tine. The male Sppatently plays some role in Tearing the young by bringing food to the lactating bitch end the pupes Pregnant and lactating fe inales require shout 300g, or 1 ines the “normal” amount, of food per day (Get, 1973) Weight at birth is about 250 to 275, ‘nd iength of the body (ip of head to hase of tall) i about {e0'mm (Grex, 1968; Bekott and Jamieson, 1975). Between birth and week 8 the average eight increase is about 0:31 kgiweck {Bekoffand Jamieson, 1975), and the pups reach adult weight at bout month 9. Eyes open st about day 14, Teeth erupt as fol- fows (on the average): upper canines (ay 14), lower canines and upper incisors (day 18) and lower incloors (day 16) (Bekoff Sand Jamieson, 1975: The young are able to urinate apd defecate ‘without maternal assistance by week 2-or 3. Jackson (1951) Scged the nesence ofthe cosing of rani wtuen, ‘ung emerge from the den in week 2 or 3.and may disperse in months 6 to 9. Not all young disperse. Although there are fo field data, consistent reports op the development of social Behavior in capive coyotes Fox and Clark, 19717 Bekoft, 1972a Wide, 19750) indicte thet young coyotes form domina lavonship via severe, unetuaized fights between 25 and 35 days ot age ‘Coyotes in captivity may live a8 long as 18 years (Young, 1951) but in wild populations few individuals ve more than to 8 years (Gler, 1908: Mathwigy 1973). Maximum ages known in the wild are’ 13.5 (Nellis and Keith, 1976) and. 18.5 yeare {Knowlton 1972) In an unexplited population, Knowlton found 70% of individuals in spring (preswhelpin) toe fess than 5 years old and less than 39% to be more than 9 years old In’autumn, gore than 80° of the individuals were less than B years eld. Knowlton reported a 40 ancual mortality rate for foytes more then Year of age, with relatively high survival Between years 4 and 8. Eetimated mortality rates in central Ak Berta (Nels and Keith, 1970) were 71% through year Land 36% 10 488 for animals more than = year old (eee table 1). In Towa, Mathwig (1973) found greatest life expeetaney at 102 years and Feast a 3M. To maintain stabity, a net survival of 33% was ‘ede in the populations that Knowlton (1972) studied, whereas cls and Kerth calculated that 389 survival was necessary Gier (1968) fet that there, were three basic limite on reproduc: on: 1) elimatic factor, 2) parasites and disease, and 9) food {Enases due to predation, accidents, and man are alvo important. ECOLOGY. More is known ahout the ecology of the coyote than perhaps any other carnivore (cee Bekolt, 1974). Many dats have been collected because of the economic interests of control and management programs, The coyote is an oppor Tunistie predator and includes wide variety of food nits dit, the percentage by volume and weight varying individually fand also with season and locality. Korsehgen (1957) listed 56 Snimal, 28 plant and six miacellaneous food hems in Missouri and A. Mure (1940) and Mathwig (1973) and others have in- fluded large numbers of plant species. Individuals in captivity 1975) require about 600g of meat per day.MAMMALIAN SPECIES 79 Fioune 6. Aggressive displays by two 23-day-old covotes Because of the varied die, itis impossible to list all food items that have been found in teats and stomachs or ebserved Iheing taken in actual kills. Deer, lk sheep, rabbite, various rodents, groundsnesting birds such as the ferruginoas hawk Angell) 1569), bobhite qual (Lehmann. 1946), and Conada geese (Vermecr, 1971), amphibians (Minckley. 1966), Izerds, Snails, fish, oristaceans, aad. Insects comprise the "meat Items. Various berries, peaches, pears, apples, persimmons, ‘watermelons, cantaloupes, and cerots ate included in the vege lables. and leather boots, tin cans and the like ate included {nthe miscellaneous ist” Overall, about 90% of their diet is ‘mammalian flesh. Tn the winter, much of the coyotes dit is made up of the carrion of lage game animals such as deer (see Sperty, 1933, ‘eho reviewed data for 10 western tates: O, Ju Muti 1995; A. Mure, 1940: Ozoga and Harger, 1966: Gier, 1968: Ogle, 1971; ‘Mathwig, 1978; Nelis und Keith, 1976) and le vegetable food is eaten. In the spring, summer, and autumn there fs an. in crease in the percentage (by volume and weight) of various rodents (A. Mutie, 1940: Nella and Keith, 1976), Fichter tal (0955) noted that's striking seasonal trend in Nebraska, was fan autumnal increase in the atization of Trt. Gipson (97D Stimmatized five studies on the food habits of coyotes in their friginal range and the compiled percentage data were a8 fllows fabbits, 1-1; rodents, 36.8 carton, 25:5 livestock, 21.9; wild Hd 1.0: der, 79: ru. 6. andl pole. 4 Most anayner of coyote predation on both large game mammals and domestic livestock indicate that young, ody and sick animals constitute the bulk of this portion of their ‘diet (A. Mure, 1044: Ozoga c a and Workman, 1913), and that coyote predation was nota primary Limiting factor on game and livestock. A Mure (1933) found that ison Hole, Wyoming, was 18522 was “neural and ony 1.49% wae “harasful” Simiaty, Mathwig (1973) found thal 5% of the covote's dict in lowa was not “harmful,” whereas 159% was “Sfemimental” Ozoga and Hanger (1960) reported that healthy deer could escape {rom coyotes and bighorn sheep are able 'o hase off coyotes (Weaver and Mensch, 1940), Davenport tal {G97 Tndicated hat most lamba loot during spring died trom fdonment, docking, und infection. Denil (19738) wrnte =. l= though fawn survival rates most probebly are closely related to ‘coyote numbers and predation, there are oer factors of perhaps rmore importance [my emphasis} exhibiting strong influence upon the growth and stability of deer herds: . There fe lite fevidence that ovate predation ie a primary Limiting factor on Dopulations of big game or domestic Ivestck. ‘Oroga and Harger (1940) wrote thatthe average chase of a deer by a coyote wan about $8 m and they observed two long fotile auempts of 432 and 4.68 km by single coyotes. Coyotes vill hunt im pairs or in larger groups (Cahalane, 1947: Dobie, 961; Young, 951: Ozoes and Hager, 1966: Gir, 1968) and will form bunting relationshupe with other predators auch as golden eagles (Engel. 1960), ravens (A. Murie, 1940), and bedgers (Dobie, T8BD. Actual coyote attacks are rarely’ observed and Tale is known about their predatory habite (White, 1973).'A presumed “coyote Kill” often has heen kelled by some other Dredator, sometimes a domestic dog (Davenport et al. 1973) Sele 1971) Tsted five criteria that ean be used to distinguish ere by capt om hve fed gn a euro’ Cayate hl may be characterized by: large patches of hide leading to Careaas; 2) separation of vertebral column in the thoracolum TABLE 2. Data on coyote movements from eight representa- tive studi distances inn) Tovenile Adult a Both Study Males Females Males Females sexes Garlough, 1940 29 Robinson and ‘Cummings, 1951 126 17.8 168 Young, 1951 36.2 Young, 1951 53 400 Robinson and Grand, 1958 45.6.2 06 Hawthorne, 1971 6.47652 Chesness, 1972 10.1 64 66 Gipson and Sealander, 1972 20.8 8.16 14 Nellis, 1975, 6 7 6 % region of adults and at the atlas of fawns; 3) nasal and maxil- ‘ribs, vertcbae, and seapulae chewed: ‘adult deer, the first extern jes Attacks ‘on the head and neck frequently ‘Ozogs and tlarger, 1986: White, 1935) a6 are belly and sump es 1871). A Shearing ite aed (Young 95) anda iteand-ea sequence {s'common, For smaler manna a stalkand-pounce approsel Is common. Coyotes generally eat the contents ofthe stomach tnd intestines of young but not of adults (Davenport eta. 1913 ‘White, 1973). See Bekott (19752, 19750) for inlormation on the Aevelment of predator bev coma pap” votes carry 2 ide variety of parantes (Young, 1951; ier, 1968; Thornton, Bel, and Reardon, 1970). Pleas are the tool common external parasitewexg. Puler simtlon, Hop Ispspllan afin and Cedtopsylla implex (rabbit Nea), Chaey. ‘psrltafororts (raccoon flea}, and Justopules porcinas Gavel fea). Other external paranies include various ticks Uozedes timulans, 1. ding and Dermacentor variable) and lice (Dem dicts) ternal parasite inclode the cestodes Tacnia preforms (Common according to Gier, 1968) and Mesocestoides corti: oundworms, Physataptera vara and Filaroides ester the trachea, bronchi, and rarely inthe lungs) intestinal worms such 3: Toncicaris leonina: dog bookworm, Ancyelostoma caninim, that can pave shrough the placentas whipwormm, Prechursvulpts heareworms, Dirfdoriaimmits: pinworms, Oxyuridae; thorn Headed worms, Oncicola canis ‘and Spriacercatupt, in the ceophagis, stomach, it coceiia fungus, Isospora Falta Coyoten also 1950: Lundgren ef al 198), abies Sie, 1972 Behm and bubonic plague ( ne In addition. eovotes suffer fom de mange (av Mure 1944, Holmes {Geamer et eft. 1068; Dietrich and an. Pelt, 1972), 0 fever {Enright ct of197), aortic ancuryeme (Thornton etl. 1979) ‘arlous cardigascular disorders (Rose and: Suruky 1973), and Serius wounds {or example, bullet wounds) missing limbs, and broken bones. Coyotes and other carnivores can live in the same area, dou copotes, like other smaller predators, do not compete: well with the wolf Cech, 1966, 1975), Coyotes generally do ‘not tolerate fores or bobcats (Young, 1951) and puma wil kil and tat coyotes (Young, 1940), - Coyotes spend a good deal of time on the move. They a active primary in ay evening, especially in winter (Grogs Sind Hlarger, 1906; Cheaness, 1912), but do show sporadic ac: tivity in the daylight hours. Gipson and Sealander (1972) showed {principal actviypeak at snaet with a minor peak at day teak Asana fn summer, they fou apimae tobe more active by day, with pups more setive by day than adults, Coyote tmovement paltcms have been atudied In various places and like the movement pattems of wolves (Mech, 1978) they can be ‘heled seemed win” “enzyme tage Aipersls, “or ae long migrations, Males tend to have larger Nome ranges than de females. In Minnesota, Chesness. and Bremicker (1973) found home ranges of mates to average 81.92 ken, whereas those of females averaged 10.08 mt. Furthermore, {he home ranges ofthe males overlapped considerably but those fof the females did not (an implication, but not proof, of ter= wal, sonal communication omalies (Nellis, 1972}, fodesta, 1968), "cance6 ritoriity). In Arkansas, Gipson and Sealander (1972) reported inate home ranges from 20.8 to 41.6 km and female home ranges from 8 to 9.6 ke. Other aties have reported home ranges of 20.8 to 32 kint Wells and Keith, 1976, 62. to OD Ke (Oxoen and Harper, 1966) and 9.6 10 128 kid (Camenzind, 1973). In Minnesota, home ranges were elongate with mean length about T8'times mean wath ‘Many data fave been cllected concerning length of move- iments of marked individuals in various locales (able 2. No eon ‘ate differences between sexes have been seen, Coyote move ments of over 160 kr are not uncommon. Oxoga and Harger (2566) found average daily travel tobe about 4.0 km. Movements of young coyotes alto have been studied (table 2, Dispersal uaaly over namin and winter (ber to ‘sbruary, some pups. not_dispersing during their fist year Dispersal occurs randomly inal diretions. Bupe wil move up- wars of 60 to 160 ke. Uafortunatly. few pups marked at dens re relocated by invenigators 22 by Nelle and keith, 1970), "The densiy of coyote populations varies with local tions. Knowlton (1973) suggested that density of 0.2 to Ok {0.5 vo Lmi® would be a realistic educated guess for densities over a large portion of the range, and hie suggestion fs well with data collected by others (oung, 1931: Orogs and Harger, 906; Giers 1968: Mathwig, 1973: and Nels and Keith, 1970) ‘Coyotes are_ usally abserved as Tone individuals (Ozoga, 1963; Oroga and’ Harger, 1960: Chesness, 1972) or a8 pairs (especialy during the breeding season) Larger grours of corotes fre probably parent) and young. Packs" of covates have been Aescribed in the literature Dobie, 196; Camensind, 197, but harmony and mutual respect (Dobie, the last 180 years thin relationship. particularly beeause of the covute's {n't recent paper by Shelton (1973), the cnvote was referred to t being a “dispensable animal." Indeed, the coyote has been Fesponsible for large economic losses to the domestic Hve- tock industry (Young, 1981; Ger, 1968: Cummings, 1972; Neese, 1918) "One of the frst bounties on coyotes was. established jin Missouri in 1625 (Young, 1950. Inthe 1050's, when beaver decreased in value, the vaiue of coyote rose to about 8.73.10 $1.50'per animal (ior more detailon prices see Young, 1951, Shd'van Wormer, 1968) Coyote fur is notin great demand (Frye Sind Vay, 1942, ranked it sath out of neven skins) and Young (G95 1:11é felt that the use of coyote skins and the success of the bounty system depended on the sGekle dictates of fashion.” The bounty tystem has not heen effective (Howard, 1973) and hat been supplemented by use of varius other control methods fh a shooting from snowmobiles (Wetmore etal 1970) and ‘iphones, trapping (ace Casto and Preanall, 1985, for a com- Parison of various trapping methods), coyategetiers.(M-#s Eyanile gun, a selective method-sce Beason, 19T). aversive onditioning’ (Gustavson ef aly 1974 Bekoth” 1975), and the ‘se of various chemieats sch as strychnine, 1680 (sue mo fuoroacetate consilered he the “best poson™=— Howard, 1973), and antierti Balser, 1964; Kennelly, 1969) Dose problems. Linhar and Kennelly (1992), developed « "marker" Eremethyehoretsacyelne) to “label” animale ‘aking baits ‘The efficiency of removing animal preyed on the tem reported damaged is also an important, ileration (Gipeon, 179). Knowlton (1972) suggested that contol Pram seul nade the removal of parca indi doa fom populations. Another control method that has worked to redlice lose of domestic livestock simply involved t Carrion by farmer in central Albert, because coyote Krcvlly on carrion apart of their winter det (Yodd and Reith, Undated). Gir (1968:25) wrote that “coyotes may be encouraged fr even taught to ill poultry and other farm animale by farmers siscardng dead anid were eoots can find then Hane T9ERGier, 195), However, ivis important to realize that few ‘eontral programe have been effective. “The lack of success af coyote control programs basically is due to the lack of objective studies ‘on the biology of the oyote and other predetors (sce Homacker, 1972, and Howard, 1979) Lite is known shout population dynamics and predatory methods. In addition, ranchers and enviroumentalissappeat ually reluctant to discuss issues (Buys, undated) Howard 8753) Summed up the problem well “The ecological re of Dredators, that of the coyote for example, ellom gets rational Eonsideration, and unproven concepts are often ‘perpetuated father than challenged sclentcaly.* The lack of patlence by those persons suffering economic losses is understansble, how fever, might prove beneficial in the end to stop using ineffective methods and harness both time and energy to more ellicient ‘scientifically based programs (Pringle, 197). Otherwise, indie: MAMMALIAN SPECIES 79 sina erp at population spceson may once ean be Wild Coyotes have been stidied primarily using raditelom ‘try and capturesrecapture methods. Both these techniques cat and clomach analyser and infreqently studied by direct sbservation, ‘The. use of trapping along with other methods provides information on age, sexs weight, size, and breeding Esndiion, and allows blood samples to be drawn. Intramuscal Injections of 25 mg of phencyclidine hydrochloride +25 mg of bromazine hydrochloride have heen set to immobilize trapped frimaie(Cheanes, 197, Consusinge frequently dan by sien Inventories and trapping” Ages af coyotes may be enimated by ‘counting dental cementom anna (inhart and Knowlton, 1967) Ulster 1979) recently suggested alternate: methods (eve lens: weight ofthe baculum, and thermal contraction of tail tendons) That appear to be as accurate ar the counting of cementum fnnoll and less expensive and less claborate BEHAVIOR. Because ofthe elusive nature ofthe covote, there hase been few direct abservations of social behaviors). Detaled stades of behavior other than those dealing with ter- ori of home range movements have heen done on captive ‘animale (Fox, 1010: Fox and Clark, 1971; Bekof, 19724, 1972), 19tla, 19730: Brown, 1973. Field observations indicate that the coyote is lens socal than either the gray wail of Une red wall {ley and'Mefirde, 1915), altough large “packs” of coyotes have been observed. Gier (1915) wrote that there is'no known focal structure other than the tantly. Young coyotes form ‘dominance relations via severe fights between 25 and 35 days fof age (igure 6: Fox and Clark, 1971: Bekotf, 19722, 19740). ‘The'elationship between early fighting and later Es tion is not clea, but there ate rankrelated behaviors that may play some Tole. For example, higher ranking animals are less Suceesefl at getting litermateao play with tiem and spend less time interacting with them (Beko 1974). These. animals to tend to remain at a greater datance from littermates and frequently are asynchronous in activity. with the est of the froup. Such higher ranking individuals may be those who Teer {eave the group. In addition, is hav also heen found (unpublished that the lowest ranking member of «liter interacts in- frequently wit litermates snd pechaps suck individuals) would als leave the group. Field testing of these hypotheses is under- way. During agonistic interactions, dominant pups and adults Spproach one another with a mitt lceged gat, care forward and rect, fur‘on the back erect (ploerectiony, the tail a about a EP tgle from the vertical and frequently snarling and exposing the teeth by vertically retracting the lips: Submission may take the form of Might, eetive avoidance, or passive or ctive sube Ileson (see Schenkel, 196%, for detailed descriptions). During passive submission the animal rollover on ne back, latens Ke carg against its head, usually retracts the lips hovigotaly bmmisive grin,” and may urinate and whine. During tive submission the animal approaches its “partner” in a low rouchowalk with the tal ether tucked or held low, and may Deriorm facelicking and face-pawing. Active: submis bly develope from food-begsing and ip have deveoged frm the porte ht the pups sse when they are stimulated to excrete by their mother or other adults (Schenkel, 1967). There are Tange increases in aggicssion atthe fime thatthe female comes into heat, Detailed dies of social play Behavior have been reported by Beko 1974a, 1975) During easy stages af courtship. the male becomes creasingly atracted 10 the females ‘urine or feces, or both (Gekoft and Diamond, 1976). See Reproduction and Ontogeny for more detail. When the female is ready to copulate she will tolerate mounting attempts by Une male and will flag her tail {o one ride. After “teing” the male steps over the beck ofthe Female and the couple remain locked a 100" for periods of 5 10 25 minutes Gores ie visual, auditory, olfactory, and probably tate signals for communication purposes, Alcorn (1846) recognizes three distinct calls equeak, howl eal, and diatres cal). Lehner {G75 through extensive analysce af coyote vocalizations, de- {ailed'I1 graded signals. Vocal the coyote is much like the Jackal (Tembrock, 1963). Visual signale such ax postures. ge ues, and facial expressions have alay been described (ox, 1970, 19150; Bekoff 19721, 19725, 1974a), The coyote appears to have a more elaborate repertoire of visual signals than do ‘ore solitary canids such ai the red fox, Vulpes oulpes, but Tess elaborate repertoire than docs the wolf (Fox, 19730). Coy ‘tes do deposit scent but the use uf this for terial demaree ‘on and identification has not been proven (see Kleiman, 1966, for alternative hypotheses). During cocial investigation there is ‘considerable snifling of various regione of the body. Whether for not scent is deposived from interdigal glands xn known:MAMMALIAN SPECIES 79 GENETICS. The coyote has 39 pairs of chromosomes (Worster and Benrschke, 1968. The autosomes are arocente oF telocentrc andthe sex chromosomes are ubmetacentric eu wl Benirchke, 1967 cited by Mech, 1974, Ferthe. hybrids Krave been produced by crossing coycies with domesthe dogs oung, 1991: Kennelly and Roberts, 1969” Silver and Siver, 1969; Siengel. 1971, with wolves (us andl upus--Voung, 1951: Kolenosky, 1571; Paradiso and Nowak, 1977; Riley and McBride, 1975), and with the jackal (Seitz, 1965) Coyote-dog hybrids show decreased fecundity Mengel, i971; Gipson ct a, 1970), See {ley and MeBride, 1975, for detailed comparisons of external characterstica of red wales, coyotes, and hybrids. “There haa been considerable controversy about the “New England anid” (cee Lawrence and Bossert, 1958, 1975 Silver and Silver, 1969: Richens and Huge, 1973), Thie canid differs from known coyote hybrids and hes incorrectly been rlerred to'as scoy-dog. The general concensus is that the New England anid is tn extreme expression of a trend already present in Gl thamnos and. cranial evidence (Lawrence and Borer, 1965} and data fom behavioral development (Bekotf etal, 1975) that New England canids predominantly are coyotes th some intogression of dog and wolf genes. REMARKS. The word “coyote” means and is taken from the Azter word “Coyoty The coyote resembles jackals in many respects and Atkins and Dillon (1971) have grouped C. latrans" with C- mesomelas (Black-backed jackal) and Ce aureus (golden lack) based on cerebellar morphology. During its’ movements, the coyote may function as a seed: carrier (Young, 1951), Kleiman and Eisenberg (1973) made de- {alled compartons of eanida and felids and Fox (1973) edited ‘volume dealing with wild ‘canids. A facly comprehensive ibliography has been compiled (Bekoff, 19745) and. Knowlton (974) also assembled « useful list of references. Valume 3 af the Coyote Research Newsletter contains abstracts of papers presented at a National Coyote Workshop (Denver, Colorado, November 1974). Many aspects of coyote biology will be dealt with in a forthcoming volume (Bekoff. 1977) would like to thank Drs, David M. Armstrong, PS. Gipson, an H Nelis, and Ronald Nowak for comments on an exer ‘drat of this report. A number of persons Kindly seat me pret ints and reprints of their oftentimes hard-to-find papers. Ms, ‘barking. dog” ‘Slater kindy typed the manuscript. LITERATURE CITED Adorjan, A. S.. and C. B. Kolenosky. 1969. A manual for identification of hairs of selected Ontario mammals. Ontario Dept. Lands Forest Res, Rep. (Wildlife) 90:1-68. Alcorn, J. "1546. "On the decoying of coyotes, Jour. Mam- ‘mal, 27:122-126, Allen, J. A. 1908,” List of mammals collected by Mr. J. H iy in New Mexico and Durango, with descriptions of new mer. Ms. Nat. Hist, 19: jes and subspecies. Bull po Aa 6, A dy, of the feria havk: ad and road behavior. 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