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itt~riality). In Arkanras. Gipson and Sealander (1972) reported male home ranees from 20.8 to 41.6 km2 and female home ranges from 8 to 9.6 km2. Other studies have reported home ranger of 20.8 to 32 km2 (Nellis and.lieith. 1976). 57.6 to 80 km2 (Ozuga and Harper. 1966) and 9.6 to 12.8 km2 (Camenz~nd. 1974). in Minnesota, home ranges were elongate with mean length about 1.8 times mean width. Many data have been collected concerning length of movements of marked individuals in various locales (table 2). No consistent differences between sexes have been seen. Coyote movements of over 160 km are not uncommon. Ozoga and Harger (1%6) found average daily travel to be about 4.0 km. Movements of young coyotes also have been studied (table 21. Dispersal usually occurs in autumn and winter (October to February). some pups not dispersing during their first year. Dispersal occurs randomly in all directions. Pups will move upwards of 80 to 160 km. Unfortunately, few pups marked at dens are relocated by investigators (22% by Nellis and Keith, 1976). The density of coyote populations varies with local conditions. Knowlton (1972) suggested that a density of 0.2 to 0.4/km2 (0.5 to 1.0Imi2) would be a realistic educated guess for densities over a large portion of the ran e. and his suggestion fits well with data collected by others (foun 1951- Ozo a and Harger, MAMMALIAN SPECIES 79 criminate attempts at population suppression may once again be resorted to. Wild coyotes have been studied primarily using radiotelem- etry and capture-recapture methods. Both these techniqurs pro- vide information on movements. Fwd hahits are inferred from scat and stomach analyses and infrequently studied hv direct observation. Tlir use of trapping along with other methad* provides information on ape. sex. weight. size. and breeding condition, and allows blood samples to be drawn. Intramusc-ular injections of 25 mg of phencyclidine hydrochloridr + 25 mg of promazine hydrochloride have been used to immobilize trap~rd animals (Chesness. 19721. Censusing is frequently done by "siren inventories and trapping." Ages of coyotes may be estimated hy counting dental cementum annuli (Linhart and Knowlton. 19671. Utsler (1974) recently suggested alternate methods (eye lens. weight of the baculum. and thermal contraction of tail tendons) that appear to be as accurate as the counting of cementum annuli, and less expensive and less elaborate. . - BEHAVIOR. Because of the elusive nature of the coyote. there have been few direct observations of social behaviorts). Detailed studies of behavior other than those dealing with ter- 1%6; Gier, 1%8: Mathwig, 1973; anf~ellii and aeith. 1976). ritorial or home range movements have been done on captive Coyotes are usually observed as lone individuals (Ozoga, animals (Fox, 1970: Fox and Clark, 1971; Bekoff. 19720. 197%. 1%3: Ozoga and Harger, 1966; Chesness. 1972) or as pairs 1974~. 1975a; Brown. 1973). Field observations indicate that the (especially during the breeding seazon). Larger groups of coyotes coyote is less social than either the gray wolf of the red wolf are probably parent(s) and young. Packs" of coyotes have been (Riley and McBride. 1975). although large "packs" of coyotes described in the literature (Dobie, 1961: Camenzind, 1974), but have been observed. Gier (1975) wrote that there is no known whether or not they operate as do wolf packs is as yet unknown. social structure other than the family. Young coyotes form The relationship of the coyote with early man was one of . dominance relations via severe fights between 25 and 35 days harmony and mutual respect (Dobie. 1%1; Gill. 1970). Within of age (figure 6; Fox and Clark. 1971; Bekoff. 1972~. 197401. the last 150 years this relationship has changed considerably. The relationship between early fighting and later social organizaparticularly because of the coyote's predatory habits. In fact. tion is not clear, but there are rank-related behaviors that may in a recent paper by Shelton (1973). the coyote was referred to play some role. For example, higher ranking animals are less as being a "dispensable animal." Indeed. the coyote has been successful at getting littermates to play with them and spend less responsible for large economic losses to the domestic live- time interacting with them (Bekoff, 1977~). These animals stock industry (Young. 1951; Gier. 1%8: Cummings. 1972: Neese. also tend to remain at a greater distance from littermates and 1973). One of the first bounties on coyotes was established frequently are asynchronous in activity with the rest of the in Missouri in 1825 (Young. 1951). In the 1860's. when beaver group. Such higher ranking individuals may be those who later decreased in value, the value of coyote rose to about S.75 to leave the group. In addition, is has also been found (unpublished $1.50 per animal (for more detail on prices see Young. 1951, data) that the lowest .ranking member of a litter interacts inand van Wormer, 1964). Coyote fur is not in great demand (Frye frequently with littermates and perhaps such individual(s) would and Lay, 1942. ranked it sixth out of seven skins) and Young also leave the group. Field testing of these hypotheses is under- (1951:118) felt that the use of coyote skins and the success qf way. During agonistic interactions, dominant pups and adults the bounty system depended on the "fickle dictates of fashion. approach one another with a stiff-legged gait, ears forward and The bounty system has not been effective (Howard, 1973) and erect. fur on the back erect (piloerection), the tail at about a has been supplemented by use of various other control methods 45" angle from the vertical. and frequently snarling and exposing such as shooting from snowmobiles (Wetmore et al.. 1970) and the teeth by vertically retracting the lips. Submission may take airplanes. trapping (see Casto and Presnall. 1944. for a com- the form of flight, active avoidance, or passive or active suhparison of various trapping methods). coyote-getters (M-44 mission (see Schenkel, 1%7, for detailed descriptions). During cyanide gun. a selective method--see Beason, 1974). aversive passive submission the animal rolls over on its back. flattens conditioning (Gustavson er al., 1974; Bekoff. 1975~). and the its ears against its head. usually retracts the Lips horizontally use of various chemicals such as strychnine, 1080 (sodium mono- into a "submissive pin," and may urinate and whine. During fluoroacetate, considered by some to be the "best poisonw- active submission the animal approaches its "partner" in a low Howard, 1973). and antifertility agents (diethystilbestrol-Baleer, crouch-walk with the tail either tucked ir held low. and may 1964: Kennelly. 1969). Dose regulation and distribution have been perform face-licking and face-pawing. Active submission probproblems. Linhart and Kennelly (1967). developed a "marker" ably develops from food-begging and passive submission appears (demethychlortetracycline) to "label" animals taking various to have developed from the posture that the pups assume when baits. The efficiency of removing animals that have actually they are stimulated to excrete by their mother or other adults preyed on the items reported damaged is also an important con- (Schenkel, 1967). There are large increases in aggression at the sideration (Gipson, 1975). Knowlton (1972) suggested that control time that the female comes into heat. Detailed studies of social rograms should consider the removal of particular individuals play behavior have been reported by Bekoff (1974~. 19756). %om populations. Another control method that has worked to During early stages of courtship, the male becomes inreduce loss of domestic Livestock simply involved the removal of creasingly attracted to the female's urine or feces, or both carrion by farmers in central Alberta, because coyotes do depend (Bekoff and Diamond. 1976). See Reproduction and Ontogeny heavily on carrion as part of their winter diet (Todd and Keith. for more detail. When the female is ready to copulate she will undated). Gier (1968:23) wrote that "coyotes may be encouraged tolerate mounting attempts by the male and will flag her tail or even taught to kill poultry and other farm animals by farmers to one side. After "tieing" the male steps over the back of the discarding dead animals where coyotes can find them." Hunter- female and the couple remain locked at 180" for periods of 5 trapper programs appear to be the most successful (Howard. to 25 minutes. 1973; Gier. 1975). However. it is important to realize that few Coyotes use visual. auditory, olfactory, and probably tactile control programs have been effective. signals for communication purposes. Alcorn (1946) recognized The lack of success of coyote control programs basically three distinct calls (squeak, howl call, and distress call). Lehner is due to the lack of objective studies on the biology of the (1975). through extensive analyses of coyote vocalizations, decoyote and other predators (see Hornocker, 1972. and Howard, tailed 11 graded signals. Vocally, the coyote is much like the 1973). Little is known about population dynamics and predatory jackal (Tembrock. 1963). Visual signals such as postures, gesmethods. In addition, ranchers and environmentalists appear tures. and facial expressions have also been described (FOX, equally reluctant to discuss issues (Buys. undated). Howard 1970, 1975~: Bekoff, 19720. 1972b. 1979). The coyote appears (1973:s) summed up the problem well: "The ecological role of to have a more elaborate repertoire of visual signals than do predators, that of the coyote for example, seldom gets rational more solitary canids such as the red fox, Vulpes vulpes. but consideration, and unproven concepts are often perpetuated a less elaborate repertoire than does the wolf (Fox, 19750). Coyrather than challenged scientifically." The lack of patience by otes do deposit scent but the use of this for territorial demarcathose persons suffering economic losses is understandable, how- tion and identification has not been proven (see Kleiman, 1966, ever, it might prove beneficial in the end to stop using ineffective for alternative hypotheses). During social investigation there is methods and harness both time and energy to more efficient, considerable sniffing of various regions of the body. Whether scientifically based programs (Pringle. 1975). Otherwise. indis- or not scent is deposited from interdigjtal glands is not known.
MAMMALIAN SPECIES 7u GENETICS. ~h~ ,.,ly~~tr. !,as 39 pain of chromosomes koff and F. E. Poirier, eds.). Garland, New.York. in press. (Wurster ant1 Renirschkr, Iwdll I'he autOsOmes are acrocentric - 1975~. Predation and aversive condrtlon~ng In coyotes. or telocentric and the c.llr,,looa"meS are submetacentric (Hsu Science 187:1096. and Benirschkc. 1967, l.il,.,~ )tr Mech. 1974). Fertile hybrids - 1975d. The communication of play intention: are play aighave been pnduced by ,.nl..i~~p .I~"YO~~S with domestic dogs nals functional? Semiotica 15:231-239. (Youn 1951: Kennellv ,,,,,I nl~l~all*. 1969; Silver and Silver, - 1977a. Mammalian dispersal and the ontogeny of individual 1969; kengel. 1971). willl wolvr. and lu u-young. 1951; behavioral phenotypes. Amer. Nat.. in press. Kolenosky. 1971: Paratil.,, Naar*, k, 1971: hey and McBride, - 19776. Coyote biology: evolution, behavior, ecolog~. and 1975). and with the jarLIll ( ~~il,,. 1wl5) Coyotedog hybrids show management. Academic Press, New York, in press. decreased fecundity (h(,np,l, IU; I ; Gipson er 41.. 1976). See Bekoff, M., and J. Diamond. 1976. Precopulatory and copu- Rilev and McBride. lux r,., .l,.la~le.d comparisons of external - - . - .... ,.,, ... latory behavior in coyotes. Jour. Mammal. 57:372-375- ~ha~acteristics of red rr,llves c.,lvl,lr~. and hybrids. Bekoff, M., H. L. Hi]], and J. B. Ntton. 1975. Behavioral There has been c.,lnriddralll,, ~~~ntroversy about the "New taxonomy in canids by discriminant function analysis. Sci- England canid" (see l.nwrencc nllll Bossert, 1969, 1975; Silver ence 190:1=1225. and Silver, 1969; Rirll,,n. ... -..- -,,A Illluir., - 1974). Fs.canid differs Bekoff. M., and R. Jamieson. 1975. Physical development in from known coyote-d,~~ lIybrid,, a,lli Ilas incorrectly been referred coyotes (Canu latrans) with a comparison to other canids. to as a coy-dog. The gr,lrral conl.cll~u~ is that the New Endand Jour. Mammal. 56:f5&692. canid is an extreme rr,,ression a trend already present m Brown. J. B. 1973. Behavioral correlates of rank in a captive C. I. thamnos and cranial evitlrllce (Lawrence and Bossert. litter of coyotes (Canis laram). Unpublished M.A. thesis, 1969) and data from khaviord drvelopment (Bekoffet 1975) Purdue Univ., Lafayette, Indiana, 151 pp. suggests that New Endand -,,ida predomnantly are COYOteS BUYS. C. J. Undated. Predator control and rancher's attitudes. with some introgression of dog wolf genes. Mimeo aphed, 24 pp. Cahalane. gf H. 1947. A deersoyote epiaode. Jour. Mammal. REMARKS. The word *'cnyote*v me_ans "barking dog'' - 28:.%99 - . - - - < . and is taken from the Aztec worcj "Coyotyl. Camenzind, F. J. 1974. Temtorial and social behavior of The coyote resembles jackalR in many respects and Atkins coyotes (Canis latrans) on the National Elk Refuge. Northand Dion (1971) have pouped C. latram with C. mcsomelas western Wyoming. Jour. Colorado Wyoming Acad. Sci. (black-backed jackal, and C. (&den jackal) based on 7:56. cerebellar morpho10~-. ~asto:~.. and C. C. Presnall. 1944. A comparison of coyote During its movements, the coyote may function as a seed- trapping methods. Jour. Wildlife Mgt. 855-70. carrier (Young, 19511. Neiman and Eisenberg (1973) made ,de- Chesness, R. A. 1972. Home ran e and territoriality of coytailed comparisons of canids and felids and Fox (197%) edited otes in northcentral Minnesota. ka er presented, 34th Mida volume dealing with wild canids. A fairly comprehensive west Fish and Wildlife Conf.. Des hoines. bibliography has been compiled (Beltoff, 19746) and Knowlton Chesness, R. A., and T. P. Bremicker. 1974. Home range, (1974) also assembled a useful list of references. Volume 3 of territoriality, and sociability of co otes in nonhcentral Minthe Coyote Research .Vewsletter contains abstracts of papers nesota. Paper presented, Coyote Kesearch Workshop. Denpresented at a Natiod Coyote Workshop (Denver. Colorado. ver. November 1974). Mac? aspects of coyote biology will be dealt ~olbe;. E. H. 1969. Evolution of the vertebrates. J. Wiley with in a forthcoming rolume (Bekoff. 19776). and Sons. New York. 535 pp. I would like to Drs. David M. Armstrong. P. S. Gipen, Cook, R. S., M. White, D. 0: Trainer, and W. C. Glazener. Carl H. Nebs, and Rmald Nowak for comments on an earher 1971. Mortality of young white-tailed deer fawns in South draft of this report. .+ number of persons kindly sent me pre- Texas. Jour. Wildlife Mgt. S47-56. prints and reprints of zheir oftentimes hard-tefind papers. Ms. Cunningham. V. D.. and R. D. Dunford. 1970. Recent coyote D. Slater kindly typed zbe manuscript. record from Florida. Quart. Jour. Florida Acad. Sci. 33: 279-280. Daniel, W. S. 1973a. Investigation of factors contributing to sub-normal fawn production and herd growth patterns. Job Adoian. A. S., and G. B. Kolenosky. 1969. A manual for Progress Rep. no. 10, Texas Parks and Wildlife Dept. the identification ~f hairs of selrccrd Ontan11 mammals. - 19736. Investigation of factors contributing to subnormal Ontario Dept. Larcr Forest Res. Rep. ~Wildlifr) 90:Id. fawn production and herd growth patterns. Job Progress Alcorn, J. R. 1%. GI the decoying of royotrr. Jour. Mam- Rep. no. 10, Texas Parks and Wildlife Dept. mal. 27:122-126. Davenport, J. W., J. E. Bowns, and J. P. Workman. 1973. Allen, J. A. 1903. 'List of mammal:. cnllrrtrd hy Mr. J. H. Assessment of sheep losses to coyotes: a problem to Utah Batty in New Meux and Durangu. with drsrriptionr nf nrw sheepmen. a concern of Utah researchers. Agric. Exp. Sta., species and subs-,ecies. Bull. Amer. Mur. \at. Hint. 19: Utah St. Univ.. 17 pp. 587-612. Dersbury. D. A. 1972. Patterns of copulatory behavior in Angell. T. 1969. -4 qudy of the fermginoun hawk: .dull and male mammals. Quart. Rev. Biol. 47:1-33. brood behavior. L-ring Birds 8:225-241. Dieterich. R. A.. and R. W. van Pelt. 1972. Juvenile osteo- Atkins, D. L.. and L. 5. Dillon. 1971. Evulu~ion a ~ thr f crrr- malacia in a coyote. Jour. Wildlife Dis. 8:146-148. bellum in the genus Canis. Jour. Mammal. 52:WeIOi. Dobie. J. F. 1961. The voice of the coyote. Univ. Nebraska Bailey. V. 1905. A budogical survey of Texan. N. .her. Fauna Press. 386 pp. 25:l-222. Dunbu. M. R. 1973. Seasonal changes in testis morphology Balser, D. S. 1964. Management of redator puhtmfi vi~b and rperamtogenesis in adult and young-of-the-year coyotes antifertility agents. Jour. Wildlife &t. 28:&. (Conu latrans). Unpublish* M.A. thesis, Oklahoma State Beasom, S. L. 1974. Selectivity of predatur contrul trchniqura Univ.. Stillwater. 24 pp. in South Texas. Ju~ur. Wildlife Mgt. 38:IUia. Eli.*. H.. and D. Schwa-. 1971. Cerebr~onical surface Behymer, D. E., F. L. ?rye, H. P. Riemann. C. E. Frmti. md aman. volumes. lengths of ' and their interdependence in J. B. Ennght. la':. Observations on thr prth~yrnrmcm of mammals. including man. y~iiugetierkunde 36:147-163. rabies: experiment% infection with a virus of cebytntr own. Illlu~t. D. C. 1W. A list of mammals collected by Edmund Jour. Wildlife Dis. 30: 197-203. HrUrr. in thr San Pedro Martir and Hanson Laguna Moun- Bekoff. M. 1972~. .a: ethological study of thr drrrlopmrnt tarn* anJ thr .ccompanying coastal regions of Lower Caliof social interactiua in the genus Canis: a dyrdlc andy~l.. futnu. Publ. Field Columbian Mus.. 2001. Ser. 33199-232. Unpublished Ph.1 . dissertation. Washimon Unlr.. St. h-1. #. L.. and f. .4. Vaughan. 1966. A coyote-golden eagle Louis. 164 pp. a.rrulm. Juur. Mammal. 47:143. 19726. The devp.dPment of social interaction. play. and 1. 8.. D. E. &h mer. C. E. Franti, V. J. Dutson, metacommunicaaun. in mammals: an etholnpical prsyr- * lehunt. M. 6. Wright. and J. E. Gaggin. 1971. tive. Quart. Rev. 301. 17:412434. Thr bhar*,r of V f:ver rikettsiae isolated from wild ani- 1973. Behaviorai fvolution in canids. Proc. Xlllth In~rrna~ mal* le~ amhrrn Cdlft~rnia. Jour. Wildlife Dis. 7:&90. Ethol. Congr., Uasamgton. D.C., p. 7. F*hhJla. )'. Zoulqischer Atlas. P 1-2. 1974~. Social pip and play-soliciting by infant ranad. t:w*f. N. b' 19fl. Thr carnivores. Cornell finiv. Press, New Yerk.Wbl, Amer. 2001. 14:32-340. -. t'whtrf. t: . Wuldman. and J. H. Spather. 1955. Some 19746. A gene- bibliography on the coyntr tCan1, IrrJby p.Iw- uf latrans). Coymar ?--pss, Boulder. 26 pp. ralyc*rr in Nebraska. Ecol. Monogr. 197Sa. - - . - - . An . - - - nvc-tpv - -. . - of social and physical dr>rlopnlrnt f!l 13. t ' J in coyotes. Coyote fLes;~kwsl. 3:12-13. fbr plan. dt* morel east. Nat. Wildlife 1.1 Ib.lb 197%. Socializanror: in mammals with an rnl~lha*l* an 10'. w 1- A rurnparatlvr study of the development non-primatee. In P=te socialization (S. ChevrLrr-Skdnb
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ROY E. DAWSON SEASONAL PARK RANGER
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historical (Simpson 1965). The obje
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qauru Ao 1 IeA lap Inog suo K qeao
Page 7 and 8:
Figure = 4 Trrnsect locations 111 1
Page 9 and 10:
Table 1. Selected study habitats (A
Page 11 and 12: ats and extirpated species from the
Page 13: Table 3 Site and Transect Locations
Page 16 and 17: faunal elements are represented by
Page 18 and 19: Onvchomv. Nwthwn Grasshopper House
Page 20 and 21: 8 House House Order ~arni vora Carn
Page 22 and 23: t mnis Jatrans Coyote Canis JUDW 6r
Page 24 and 25: ermwhi lus ) rtcrrl i s 601 den-man
Page 26 and 27: .Order Insectivwa panus Dwarf Shrew
Page 28 and 29: Order Insectivora WNDEROSA PINE cin
Page 30 and 31: individuals were caught in pitf a1
Page 32 and 33: REFERENCES Armstrong, Dm PI. 1972.
Page 34 and 35: Order Marsupi carnivora I POTENTIAL
Page 36 and 37: t h r m ncarl oti; Uestern Harvest
Page 38 and 39: SPECIES ACCOUNTS OF OBSERVED &NI MA
Page 40 and 41: FIGURE 2. Skull and jaw of Zapw hud
Page 42 and 43: 0 a douthitti, to be more abundant
Page 44 and 45: and 22 October, 1 November (2), and
Page 46 and 47: CryptOtis parva. By John 0. Whitder
Page 48 and 49: MAM!tIALIAN SI'ECIES NO. 43 abscnt.
Page 50 and 51: MAMMALIAN SPECIES NO. 43 and Wharto
Page 52 and 53: MAMMALIAN SPECIES NO. 43 7 Dryden,
Page 54 and 55: MAMMALIAN SPECIES No.'56. pp. 1-3.
Page 56 and 57: MAMMALIAN SPECIES NO. 56 / Known he
Page 58 and 59: 2 MAMMALIAN SPECIES 79 LUPUS pambas
Page 60 and 61: a &=station 4 MAMMALIAN SPECIES 79
Page 64 and 65: a Horn. of facial expressions in ca
Page 66 and 67: Spermophilus tridecemlineatus. By D
Page 68 and 69: MAMMALIAN SPECIES 103 3 ing emergen
Page 70 and 71: MAMMALIAN SPECIES 103 5 ground squi
Page 72 and 73: 2 FIGURE 1. Skull of Sylvilagus aud
Page 74 and 75: LITERATURE CITED Allen, J. A. 1877.
Page 76 and 77: FIGURE 2. Skull and mandible of Syl
Page 78 and 79: 4 MAMMALIAN SPECIES 136 1939). The
Page 80 and 81: - 1904. Mammals from Southern Mexic
Page 82 and 83: a Pelton, 8 MAMMALIAN SPECIES 136 -
Page 84 and 85: 2 MAMMALIAN SPECIES 159 FIGURE 1. D
Page 86 and 87: a and a dent 4 MAMMALIAN SPECIES 15
Page 88 and 89: Q f Microtus prnnsvlvanicus) as a l
Page 90 and 91: 5 8 MAMMALIAN SPECIES 159 ! eters o
Page 92 and 93: 2 MAMMALIAN SPECIES 167 FIGURE 2. D
Page 94 and 95: MAMMALIAN SPECIES 167 Coulombe. H.
Page 96 and 97: Odocoileus hemionus. 'B~ Allen E. A
Page 98 and 99: MAMMALIAN SPECIES 219 responsible f
Page 100 and 101: MAMMALIAN SPECIES 219 captive 0. h.
Page 102 and 103: MAMMALIAN SPECIES 219 Anderson. A.
Page 105 and 106: Neotoma mexicana. e, John E. Comely
Page 107 and 108: MAMMALIAN SPECIES 262 The baculum o
Page 109 and 110: MAMMALIAN SPECIES 262 during severa
Page 111 and 112: MAMMALIAN SPECIES 262 MURRAY. K. F.
Page 113 and 114:
L 10 mrn FIG. 2. Dorsal, ventral, a
Page 115 and 116:
4 MAMMALIAN SPECIES 272 the facial
Page 117 and 118:
direction of the danger stimulus; w
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8 MAMMALIAN SPECIES 272 LAYNE. J. N
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EASTERN COTTONTAIL -Sy lvilaws f lo
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--39- DESERT COTTONTAIL Sylvilagus
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-- 48-- THIRTZEN-LINED GmUND SQUIRR
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BLACK-TAILED PRAIRIE DOG Cynomys lu
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Thomomy s talpoides Distribution.-T
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WESTERN EARVEST MOUSE 'Reithrodonto
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--69-- MEXICAN WOODRAT ~istribution
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-- 76- PRAIRIE VOLE Microtus ochrog
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COYOTE Canis latrans ~istributian--
Page 139 and 140:
Odocoileus hemionus Distribution--T
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Common Name Scientific Name Habitat
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