University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln USDA National Wildlife Research Center - Staff Publications U.S. Department of Agriculture: Animal and Plant Health Inspection Service December 2003 Coyote (Canis latrans) Marc Bekoff Eric M. Gese USDA, eric.gese@usu.edu Follow this and additional works at: http://digitalcommons.unl.edu/icwdm_usdanwrc Part of the Environmental Sciences Commons Bekoff, Marc and Gese, Eric M., "Coyote (Canis latrans)" (2003). USDA National Wildlife Research Center - Staff Publications. 224. http://digitalcommons.unl.edu/icwdm_usdanwrc/224 This Article is brought to you for free and open access by the U.S. Department of Agriculture: Animal and Plant Health Inspection Service at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in USDA National Wildlife Research Center - Staff Publications by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln.
Bekoff, bl.. and E. M. Gae. 2003. Coyote (Canzs inmons). Pqa 467451 in Wild ~~ of Xorth.lmaica: Biola~, Mmqacnt and Cm-Uoq 2d doon Edrred by G. A Feidhamer B C. Thompson and J 4 Chapman. Johns Hopkrns Ihivenir). Press. Bakimure, Llqlund Coyote Cunls latram Marc Bekoff Eric M. Gese NOMENCLATURE ~omov SAKES. Coyote, brush wolf, prairie waif, 9e.d woif, Steppen- &f. iobu,.hencm jackal; the word coyote means 'barking dog" and,,&en tiom the.\rrec word coyor1 ~cmmic Nhm. Canu lnrrans The coyotc is one of eight recoqizsd species in the genus Cmis. By &e late Pliocene, the ancesmi coyote, Cdnzs l<~ophnps, was swldespread in Sanh.America. The Eastem coyote, formerly reimed to 1s me 3ew Enziand cmld appears :o be a recent immigat, having prelomlnantly coyote ancesny with samc innogression of woif (C. lupus) anddag (C./Lmilinris) genes (Lawrence andbossert 1969, 1975; Silver md Silver!969; Bekoff et al. 1975; Hilton 1975; Wayne and Lehman 19921. Lehan s: al. (1991) and Wh~e md Lzhman (1992) presented cvidence that coyotes have interbred with wolves in arms,where wolves arc m e ma conspec5c mates may not be re~dily ava~lable. Subspecies. There ue 19 recognized subspecies of C. liltrum. Howzwr. because of the mob~liw of covores, the mteqnty of individual scbspecies and :heir monotic uhh& are quest~on;ible(liowak 1978). In chronolo~cal - order ofheino. name4 the subspecies are C 1. lurram. C 1. ochmpus, C. 1. co~ortis. C 1.fNsrmr C 1. lesre;, C 1, meumi, C. I. microdon. C. l.peninsu1oe C 1. vi~'lis, C 1. clrpfim, C 1. impavidm, C. 1, goidmani, C 1. rerenrls, C. 1. jnmrsi, C 1. dickeyi, C. 1. incolahu, C. I. honaurensis, C. I, thornnos, and C 1. umpqurnsls (Jackson 1951). Frcm 22.1. Dismbution o ihr coyote!,cnnls iofram). Coyotes are Nearctic canids. They occupy many dive:se habitars between about IO"N latitude (Costa Rica) and 70"N latitude (norrhem.uaska). X~ey are found hubout the continend United States and in many areas of Canada (Fig. 22.1). In some southeastern states, suchas Clonda (Cunningham md Dunford 19701 and Georgia (,Fisher 1973, and sevexi emtern aaies, it appexs that coyotes wcre wanspianted lnuoduced or :ibcr;ired by humm (Schuitz 1955; Hill ct dl. 1987). See kloorc and Pxker (1992) br the hstory of coyote expansion md changes m dismbution. With ns abiliry to adapt to iill emonment mudified by humans, coyores are now obsexed in large cines (Shargo 1988; Qumn 1997% 199%; Gnnder md L-ausman 1998: F&el 1999). Coyores xe often coniused..with other ;mias, such as -my wolves, red wolves (C njls), and dornesnc dogs. Coyotes can successiully interbreed and produce fenile hybxds wirh dl these 3prcles (Dice 1942: Ksmelly md Robcm 1969; Kolenosky 1971). Howeve:, coyotes cx Uually be afierentiated (although overla? andhybddization can ucm: 'd~~lg serolo~c pume:ers, ienral ~.i.xac:erisncs. cmal measuremezts. ne-oanaromcai ie~nues. diame:er of the nose?ad diame:er 'frhe 'hind:oat?ad, :ar!engh, mck size. 3?r.ue irnph.?elqe. behav- :or. mo :ezrccs ;for re\ie.xs. see L~wrenc; md Bossen 1967: Bekoff 19-. Eider. and &yarn 1977>. For exampie, coyotes xe typically ' 3 % smaller than gray wolves (Table 22.11, thus the nose pad (about 25 nun in diameter) and hind foot pads (less than 32 mi are cor;espondiigiy smaller. Coyotes may be differentiated from dogs wig the ratio ofpalatal wdtb (distance beween the her mar& of the dveoii ofthe upper iirst molarsi to rhe leu& ofthe uppermolar toothrow (from the anterior mxgn of the aiveolus.>f the 5131 premolar to the posterior margin of the idst molar alveoiusj (Howard 1949). If ihe toothrow is 3.1 times the palad width, the spechen is a coyote; lf the raho is less than 2.7, the specimen is a dog (this method is about 95% reliable). Tie coyote has a relafively lager braincase than C 1u.pu.s Ovlech 1974). The coyote brain is anatomically different from that of gray wolves iradins!q 1973;.4tkms 1978). The wolf has a dimple in the middle of rhe coronal,ws, whexeas the coyote does not (see also.%dans and Dillon 1971). Tnere is no wedap when comparing large coyotes :o small wolves ln zygomatic breadth (greatest distance across qgomarai, gestest length of the skull, or bite raoo (wdth across the outer edges of the alveoli oithe mrerior!obes ofthr uppx camassials divided by the length oithe upper rnolar:oothrow) (Pmdiso andxowak 1971'1. C larronr is usually smaller than C.?Em and there is almos1 no ovenap between them m gestest length ai the skull. Red wolves ~iso have.: =ore pronounczd sadrtal crest -h coyotes.?iiuitivanate ;e:iruques :we cleariy shorn coyotes, wolves, and dogs can be dizerennated anaromcally!lawrence and Bosserr 1967; Eider md Bayden 197;) and behawiomlly (Bekoffer al. 1975; BekoE1978) and 467
TABLE 22.: Representanve mean coyote we~ghts kom a vanety of locales Add* &g) Juveniles Cxg: Sowe,Males Females Xaier Eeades Sf~*Wince Gier 1968 14.1 11.8 - - Kansas Hamhame 1971 11.2 9.8 M Cmomja Richens and Hqe 1974 15.8 13.7 - Mame - Andrean and Boggess 1978 13.4 11.4 - - Iowa Berg and Chesness 1978 12-13 11-I? l&ll 10 Mhmesara Boggess and Henderson 1978 13.1 11.0 - - Kaasas Bmn 1978 13.1 11.5 - - Aikm L~Gtis 1978 14.7 12.1 - - Oklahoma,Umy md Bouim 1991 10.3 8.0 - - Yukoo Thurber and Peterson 1991 12.9 11.1 - - Alaska Windberg et al. 1991 10.6-11.4 9.1-9.6 10.&10.8 8.G8.9 Te~as - Podle et al. 1995 12.5-16.0 11.&14.2 - Quebec Windberg 1995 10.9-11.0-8.8-9.2 8.0-8.4 Texas provide for more rigorous analyses than do univariate methods. Further rehement of genetic techsues will also assist in differentiation of thr canids (e.g; Lehman et a i 1991; Wilson et al. 2030). Size and Weight. Covotes are about 1-1.S m in bodv lenpth; the tail is about 400 --long. size varies with geographic lockle &d subspecies (Jackson 1951; Hall and Kelson 1959). Adult males are usually heavier and largcthan adult females (Table 22.1). Temporal changes in coyote momholommav. -.. be occunine insomeparts ofnorth America ( Sche and Lavlgne 1987, ~hurberand Peterson 1991), but see Lanwkre and CrEte (1993) and Peterson and Thurber (1993) for rev~ews and commenls. Pelage. The banded name of coyote hair is responsible for the appearance of the blended color, giay mixd wjth a reddish rint. Coyotes show great -miation in color, ranging from airnost pure gray to IU~OUS. Melanistic coyotes are rare (Young 1951; Van Wormer 1964; Gipson 1976; Mahan 1978). Texhlre and color of the fur also vary geographically In northern subspecies, the hair is longer and coarser. In desert habitats. coyotes tend to be fulvous, whereas those at higher latitudes are more gray and black (Jackson 1951). The belly and throat are paler than the rest of the body. Course guard hairs are about 5&90 mm long; in the mane, they tend to be 80-1 10 mm. The!he underfur (up to 50 mm iongj has coronal-shaped cuticular scales (Adojan and KolenosA~ 1969; O ~le and Famis 1973). The summer coat is shoner than the winter coat. Coyote haumay be differentiatedfrom hair ofdogs and red foxes (Vuipes wipes) by the nurnbez, odder, and color of the bands, the cross-sectional mmslucence and shape, and the coronal scale pattern (Hilton and Kutscha 1978). Coyote hairs 9ically are coarser, longer, larger in diameter, and rougher and stiffer The coyote's fur is similar in insulative value to that of the,my wolf (Ogle and Fams 1973). The critical temperature of C. lavans is -10 C (Shield 1972). When wearhg the shoner summer coat, there is a decrease of about 87% in thermal conduaivlty (Ogle and Farris!973). There is usually one main molt between late spnng and an-. About 50 mm down from the base of the tail there is an oval rail gland (JIiidebrand 1952). Skull and Dentition. Among adult coyotes, males show greater development of the sagtttai crest than females. The dental formula is I 313. C 111, P 414..M 2!3 (Fig. 22.2). The skull of a mature male is about 180-205 mm long from the tip of the premaxilla to the posterior rim of the coronal cres (Gier 1968) and weighs between 170 and 210 g. PHYSIOLOGY Central Nervous Svstem. Althou& the cerebrum and cerebellum With respea to cerebellar morphology, coyotes may be distinguished fiom all other canids as follows: The anterior lobe is more than one half the rota1 width. the pardoccular process is relatively prominen\ the vermian lobule reaches its greatest size, there are fewer and larger posterior hemispheric foiia, the posterior ventral pard3occular limb is reduced in size, and there is a broad vermian hvist (Atkins 1978). The remainder of the cennal nervous system, the brain stem and spinal cord is similar to that of the domestic dog. Adrenals. Coyote adrenals are similar m smchlre to those of most other canids (Heinrich 1972). In males and females, the lefi adrenal is heavier than the right, and the adrenals of females tend to be heavier than those of males. Audition and Vision. The region of maximal sensitivity to auditory shmuii is 10&30.000 Hzwith alimit ofa~~roxlmatelv 80 kfiz Petersen et al. 1969). The retina is duplex and hbs a prepdnderance'of rods. The absolute scotopic (rod) threshold is about 1.4 fi-candles and the adaptation curve shows distinct rokone b~&s (Horn and Lehei 1975). REPRODUCTION Genetics and Hybridization. The coyotehas 38 paus ofchromosomes (Wurster and Benkchke 1968). The autosomes are acrocenmc or teiocenmc and the sex chmmos&mes are submetlcenmc (Mech 1974). Fertile hybrids have been produced by matings of coyotes with domestic dogs (Dice 1942: Young 1951; Kennelly and Robexts 1969; Silver and Silver 1969; Mengel 1971), red and gray wolves (Young 1951; Kolenosky 1971; Paradiso andnowak 1971; Mey andmcbride 1975). and goldenjackals (C. aureus; Seie 1965). Cayotedog hybrids exhibit decreased fecundty (Mengel 1971; Gipson et al. 1975). Hybridization between coyotes and red wolves is becoming problematic for red wolf recovery in the southeast United States. Anatomy and Phvsiology. -. There are no detailed reporrs of the gross or mcroscipic anatomy of coyote reproductme systems. Patterns arepro ductive hormones and reproductive behaviors in coyotes have been de- reproductive anatomy andpbysiology, there appearto be only -or differences., if am... between covotes and domestic dogs (Kennelly 1978). Berg and Chesness (1978) found no correlation be&eencarcass we~ght and ovarian weight (n = 105, r =,218,~ >.05). Kennelly (1971, 1978) documented spermatogenesis and the estrous qcie. Proestrus lasrs about 2-3 months and e m up to 10 dqvs. depending on locale (Hamlet? 1939; KenneUy 1978). Copulanon ends with the copularory Yie." during which h e (up to 25 min) the male's penis is locked in the female's va- (Grandage 1972). Juvenile males and fenales are able to breed (see below), although juvenile females may date less than adult females.
.J..],1 L~:-.-,..,re naunr:ag itrsrr.co by Xr mde ;nu ~ :il -la2 her-ali :c one j:dc &%en %~n.g. ;he male rreps ovsr the krmie's 5acx an11 &e sauple reaalr. lccksd it :SO' for ip io 25 7112. ihi same?ax nay ired iom ye= :o year, bur nor neitssaciy for!ze. ;\iui! csyores may nmw p a boc& and \wheip Jr slre pups even wnen!&i2 yem oiage igese i990). Pregnancy Rare. Cestntion. and Litter Size. Tbe?er;:suge of ;emales dia bred m a gvex :,exu.v&<es wt;h :ocai cmdidons (Gier i 968; FL?owlton 19-2; Gipsan :? i. '9775: Gese 6r xi.!989a; K?owlton md Gese 1995). Fuod slipply :s usudly :he prime izi:or; in :ood yeus. nore females, especially jsxkgs. breed (Cie: '.968; Kilowiton ma Gssc :995). Uslliily, about 6&90!6 oiaddt :emales md &:09'i or' female year!ings.~:ll produce litters @howlton 1972; Gese et a]. 1989a;.Knowi;on st 21. 1949). Tne gestest mual v,r;ation ;n ;he number cibrecding females is :e!ated to the number oijuveailes that become sexuai;:, mature (Xnowlton 1972; Kexelly 1175; Gese c: il. 1989a). ;\duits may show mois placental scars &an yr~:ings: however, Neilis md Kt:& (19761:eported that the diffcrsnce was nor itatisticaily sigruficanr. Gicr (1975) esrhated rhat the number ofyoung born was about 80% ofthe ova shed and Knowlton (1972) esrimated that about 87% oiimpiants wcr: represented by viable young. Gipson et al. (1975) reportrd ti:e =can number of ova per breeding female was 6.2, with 4.5 (T?') becoming irnpimrcd. Hanlert (1939) and.bdei1(!96.1) reported that 85-92?4 of cmbvos de.ielop into viabie young. Gestationlasts approxlmarely 63 days. Avemgelitter slze is 6, but it :s known that liite: size is affected by population ds::siry and food mailablliry the previous w!mer (Xnowlton 1972: Gese <: al. i996a. 1996b; Knowlton st 31. 1999). Knowiton (1972) reported avenge liner sizes of J.3 at h19.i popuianon densities and 6.9 at low densiiies. Gier (1968) re;rorted tie ~Eicts of hod on litter size. Duing ;~exs oihgh rodent dtnsir): mem liner size is hi&er (S.84.2: than h years ui reduced densities of rodents (44-5 1). In nonhen? iiimcs, coyote litte: size c5anges in :espanse to cycles of sno\vshoc hares (Lepus omericunus) (Todd it a]. 1981; Todd and Keith 1933; C3Donoghue e: l. 1997). Gcs; er al. (1996a) hund an increxse h coyote lice: sue afier coid snowy winters ic Yellowstone Yanonai Park. Wiiren wtb harsh temperatures and lie? snow incre3srd!he number of mgu1;lte carcasses (because of soy016 predation and ~~eather) available to odatmg females. The sex ratio oflittcrs is generally i:l, though it may be skewed toward females in ares oibigh ~xploiration &owlton 1972; Berg and Chcsness 1973; Keiman and Brady 1978; Gese et d. 1989ai. F:c- 12.3. SMI of the coyote (Cmms lirianr). From tap ro bunom: laced eew oicrmum i~rcnl mew oimanciiblc. hnd view aicmum vclltni vie;" oicmm. dorsal new aimandr'nle. blaies and females show annual q~cii changes m rqroductive anatomy =d phys:ology (Kennelly 1978). Females ue seasonally monestrus, showing one period of '%~JI" per year, usdly dunng January and blxch. depending on ~eogmphic loc~le (Hamien 1939: Gier 1968; Kcnneily 1978). P3iT Bonding. Tae dynamics of hete~osexd pair bonding are not known ior d d cayores in any detal. Data on cqtive indinduals do not appru to &Err sipficantly Erom those forw4dcogores. Caunship nay be~n as long as 3-3 mocihs before succzssiui copulation 1Bekoff and Diamond 1976)..&soc~ated c:iar,gm in behavior iave been <ercn'ccd esoeaalb inczezes m iccnr nar:mg and bowling ubsemed ar me bermmg of~.c breedins rmsm ibekotyau31amona :9-6: Wsils md a&d 198:: ECC~~S!ZOO: GCSC ma RUI? 1%- :%x). ~ur=y exiv. surirs. 31 comsaip. -he maie ';cc3mes lnc1~3~in'~i:i ~mc:ed :o he i'cmje's m e and feczs. RJcn he femaie is re3dy ro copulate, she ;o :he base ofthe tii1 is 2bour 160 mm (Gier 1968: ~ekoft&d ~amiesan 1975).Eyes opeontabout 14 days. Tetherupton thesverxyc as follows: upper curs, day 14; lower c~xnes ind upper incison, day 15: md lower incisors, day 16 (Bekarl and Jmeson 197S). Tnz young are xbiu to wnnte and defecate on theu own by 2-3 weeks. They emerse from xhc dm at about 3 weeks. Ycunq arc cared for by the mother and by other "helpers,' usdly siblings fram s prcxous yex (Brkoorf and Wells 1980: Hatier 1995). Food ~rovisioning ofthe female may occm dunng ihe nursmg period (~arkr 1995). 6 c lpha male md orbrr ass~ciates will dso help to rex the young by providing food as the young gow older (Hatier 1995). Young are ~eaned at about 5-7 wee.h (Snow 1967). Tney be~m to sat solid food at about 3 weeks. when the caregvers reprgmte semisolid ibod. Beween bmh and week 8, averaxe weight increase is about 310 stweek. Tne pups reach adldt we:ght ar about 9 months. ECOLOGY >lore :s know about he ecolog oi ;oyotrs -Sar.?erh~s any other carnivore IBeioiT 2001~). Co"otes ocxpy a vw,,ey of ilabiuts. inoicding g3ss;ancs. iescns, ma.mountam.?ills? lo not icmpi:e Well ~,rh iqc: cjriuvorcs xi xay be hilcd by them or ivo~d areas ad hablraz >cc~~.ed hy these s?ec:rs. Studies $ax documenred direct md ndoke~: -ompencon with larger cmlvares, mch 3s waives (Mech
1966,1974; Kreftmg 1969:Fuile:andKelth 1981: Tnurberetal. 1991: Peterson 1995:.Qo and Pierschrr 1999: Crabtree and Sheldon 1999) and cougan, Puma concolor rioung 1951; Boyd and C'Gm 1985; Koehler andhomocker 1991:,Murphy 1998)..4qoandP!erscher (1999) documented behamoral chanses in coyotes after several years oi m- cremed wolf abunhce innorthwestem Montana. Paquet (1991. 1992) reported that coyotes did not spaualiy avoid waives, bur actually followed their hails and scavenged woif-killed ungulates. Inten~eciiic killing amem.. to be common m carnivore communities (Peterson 1995: Palomares and Caro 1999). In Yellowstone National Park, Crabtrez and Sheldon (1999) ;:ported that wolves killing coyotes during the umters of 1997 and 1998 reduced coyote numbers: average ~ack size decreased!?om 6covotes to4 ia33% aechei. However. Gese et al. (1996% 1996b) documented simiar annual vanations m coyate pack sies in the same area in the absence of wolves. Average pack size changed from 4.6 to 6.8 coyotes during tbree winters (199&93) seven1 vears before wolf reinuoduction. an increase of 32%. Thus. annual variation in coyote pack sue and population size amibuted to wolf reinnoduction should be viewed in the context of baseline population data that documented these same fluctuations had occurred before wolf reintroduction. Changes in the abundance of food resources, mdy cyclic lagomorph populations, causes even greater muai vaxiations (3-10 times) in coyore populations than populations exposed to wolves (Todd et al. 1981; Knowlton and Stoddart 1992; O'Donoghue et d. 1997). Duect predation and competition for food and space with wolves may limit coyote numbers in some areas and under iemin environmental conditions (Peterson 1995; Ajo and PletscSer 1999). Coyotes may not to1e;ate red foxes in some areas (Voiy and Earle 1983; Major and Sherbume 1987: Sargeant e? d. 1987: Harrison et al. 1989; ~&~eant and Men 1989), butappear to be mare tolerant of red foxes when food a abundant (Gese et d. 1996d). Coyotes also benveen coyotes &d these small canids is typically mediated by resource partitioning (Rails and W'ite 1995; m te et al. 1995; Cypher and Spencer 1998; Kitchen et d. 1999). Bobcats (Lynr ru/us) also may not be well tolcrated by coyotes(yaung 195l), but Major and Sherbme (1987) foundno evidence of interference competirion between bobcats and coyotes. Soule e? al. (1988) and Crooks and Soul= (1999) reported that coyotes in southern California apparently connol the abundance and dismhution of smaller predators. In the absence of coyotes, these mesopredators (i.e., foxes and feral cats) increzse m densit). They prey on narive bird species and negatively impact the avifaunal cornmunit/ Henke and Bryant (1999) documented that coyotes were considered a keystone predator shaping faunal community smcture in west Texas. Population Regulation. Popuiation demopphics of coyotes have been studied throughout North America (e.g., Gier 1968; Knowlton 1972; Todd andkeith 1983; Windherg 1995). They exhih~taland-tenure system of exclusive territories (Camenrind 1978; Bowen 1981, 1982; Messier and Barrette 1982; Windberg and Knowlton 1988; Knowlton and Gese 1995), and with resident packs. they display a dominance hierarchy similar to that of wolves (Camenrind 1978; Bowen 1978; Bekoff and Web 1986; Gese et a]. 1996a). Tne social orgamization and land-tenure system mediate the re~lat~on aicoyote numbers as packs space themselves across the landscape in relation to availabie food and habitat (Knowlwn and Stoddart 1983; Bekoff and Welis 1986; Gese er al. 1988% Knowiton and Gese 1995; Knowlton et al. 1999). The social hierarchy and dominance smcrure among members of resident packs also iniluence accessibility to food resources (Gese e: d. 1996a. 1996%). Older. expenencedpack members are more succzssfd hunters of lar_ee prey (Gese and Grothe l995), have geater access to?logdate carcasses (Gese et al. 1996b), and are more proficient hunters of small mammals (Gese 2t a!. 1996~). Trarsienr or nomadic coyotes aiso exist across the landsczpe (Camenzmc 1978; Bowen 1982 BekoiT and We!ls i980: Fese e! al. 1988a) and may nove in10 terncones wheneve; vacancies occur. T.am 22.2. Coyote densrties in different geoppkic areas and seasocs Source Glcr :968 Knowitan 19iZ Chesness and Brcrmcker 1974 Neh and Kath 19i6 B-n 1978 Todd et al. 1981 Pvrah 1984 ~.- Gesc et al.!989a Babb andkmedy 1989 O'Donagbue et ai. 1997 Henke and Brvant 1999 Locsnon Denriv (hdi~duahh') Kansas 0.8* Texa3 0.9': :.5-2.3c.kesora 0.24.AL Ubem 0.146' AIbRta 0.46'; O X d Alberta 0.081)qdc Montana 0.15'; 0.39' -.exas 0.99 Colorado 0.26-0.338 Tennessee 0.358 Yukon 0.01-0.09F Texas 0.124.14~ Coyote densiry varies geographically and seasonally (Table 22.3) in response to changing foodresources. Available food whether rabb~ts, rodents, or ungulates, is the major factor regulating coyote populations (Gier 1968; Clark 1972; Knowlton and Stoddart 1992; O'Donoghue et al. 1995). Numuonal pressures for hmited food resources are mediated'hroueh social dominance and territorialihi (Knizht 1978; Davison 1980; ~niwlton and Stoddart 1983; Gese et ;I. 1985a; Knowiton and Gese 1995, Wmdber~ 1995, Knowlton ct d. 1999). Food abundance Todd etal. i981; Todd &d Keith-1983; ~ idd 1985; Brkoff and Wells 1986; Mills and Knowlton 1991; Harrison 1992; Windberg 1995; Gese et d. 1996% O'Donoghue et al. 1997; Knowlton et al. 1999). In areas with hard winters, when carrion biomass is low, coyote pack sizes remain small and the core social unit subsists on small mammals (Gese et d. 1996a, 1996h). In contrast during winters when carrion biomass is gcater, more coyotes remain in their social groups and pack size increases (Gese et al. 1996% 1996b). In additioq domant individuals in resident packs have greater access to carcasses and are thereby less likely to disperse (Gese et al. 1996% 1996b). Thus, dominance plays an important role in regulating coyote numbers. Knowlton et al. (I 999) reported that the acquisition of a temtory is also important because resident coyotes are more apt to survive, have more breeding oppomnities, and are more likely to have access to carcasses in winter than are nansient individuals (.4ndelt 1985; Bekoff and Wells 1986; Gese et al. 1989% 1996a 1996b). Withln resident coyote packs, size and structure change seasonally (Xnowlton 1972; DavLFon 1980; Bekoff and Wells 1986; Gese et al. 1996a). Pack slze increases during the whelping season (April), followed bv a gad& decline as pups die or dsperse and associated pack membersdisperse dming as food re&urces become more limited @howlton 1972: Davison 1980; Gese et al. 1996a; Knowlton et al. 1999). If more food resources are available over wmter, pack size may mcrease (Gese et al. 1996a, 1996b). Effecn of Exploitation. There have been few studies of unexploited coyote populations (Andeb 1985; Bekoff and Wells 1986; Crabtree 1988; Gese et al. 1989% 1996% Wlndbeig 1995). These populations generally diffcr!?om expioitedpopuiations byhaving anolder age smc- we. h~gher sumival rates, lower reproduct~ve rates, her pack sue, and!owerrecruimentlnw the adultpopulation i.4ndelt 1985; Wmdm et al. 1985; Gese eta1 1989% 1996% 1996b: Widberg 1995; Knowiton et d. 1999). Under hig levels of exploiration coyote populations
,.ailv have 1 :iol;r.ge: 3 % ~ scdc?r:. lower sum:yai,,hc:rasej num- 3,:;,~i:iczling3 rs?rciucsg, mcressd liner %re, and reiacve!y smdl,ac& (Cie: 1968: :L~owiron 19-2 Berg and C:kcsness 19-3: Cav:son 1980:.hdeit :987; :uowlton st 11. 120C). L;ne; rze 2.y mc:ese due rc reduced coyor; ;ensir,,!ik;iy n respomc ro reducrd compt.5- tion for food (,bde!r 1987. 1096),,r brerdir.3 song younger females (jlmw-lton er d. 1999). pie E~xpretmcy. Cayares J: c3pr:vity may iivc as long u I8 :,ears piou~g i951!, bur 2 wtia?opuianons. 5fe sx?ec*imcy is considerably shorter. >I&vaum 1Zes re?oned fcr wid soyctss ae lj.5 (Xsiiis in6 %:th 1976), 14.5 (IGowitcn l9i3), and 15.5 yeas (Gese 19%)). Dens. Cayotes den in a vxetl, of placcs. iccluding brush-covered siopes, s:erp jar&. under rocx ledges. thicks:~, md hollow lcgs. Dcns oiother m,als. ulciudins badgers : Toxiden ims I, airc fiequentlyused. Eem mzy have more!ha one enmnce (Bmison and Gliben 1985) ad ohen there are many hrercamec:in~runnels. Dcns may be osenred :o 'he iourh :o mar.jrize solar radiation igier 1968; Hallen 1977; Harnsan and Ci?om 1982). The sme den :nay be used iiom yrx to year Dm iharaf occurs oniy rs;.ely (?jeiiis and Kdith 1976; Cmznzlnd 1978). >Iavement of pups &Tong dezs is vey cumon (Hamson and Gijben 198:). Rasons ior these moves nc nor known. but disrurbance (Harrison 3nd Giiben 1985) ind possibly iniesration by parasires may he hcrors. Most moves are rtia"vely shar;; however, moves oi >? hi are common iharnson md Gilben 1985). \in Wormer I 1964) reponed thaf a maie coyote mcved four pups, ir.dividually, a distance of 8.3 :a. T?x den and pup-rear,rz icn'rrties are the focal joint for coyore Pamilies far several months untd puns are iaizr and ;r.obzle,.hdelt e: al. 1979; 5zr;:son a d Gilben 1985; Bekodand Wells 1986; Haticr 1995). mnvity and Movements. Coyote:; xz acrive rhroushout the ;iv. but rend io be mure :ic:ivc dunng the sariy mornlng and iround sunset (WoodmE mu Keiler l?;i2:.r\ndrit 1985; Ccse er 31. 1989b). Gipson imxi Sealmdcr (1 9-: snowed a 9racrpal acrlvity pc~k at sunset and a mor peak at kaybicsk in.arkansas. Activity patterns may chmy sessonally (Hobman ei ai. l9923j or in response to human disrurbac: and persecution (Grse et al. i984b; Kitcirn et 11. 2000a). Sczonal ac:iv??, Fatterns dso are obvious, sspeciaily dunng wmer months, when there 1s a changeb food bae (Bskoffand Wells 1936: Gese et al. 1996a). For example, coyotes living in nonhen climates rest morz dmlng winter months, when they are drpendcnt pnmanly on ungulate canon fur food thandung other se~sdns, when they fe'eedmsinly on smali rodents (Bckoh~nd Wells I98 1. Gese et ai. 1996ai. Hunrmg artemprs also decrease durn5 winrer monks, when rodents are relatively inaccessible brxuse of snow-covered pun4 one of &r mijor rodent food item (Linta yround si;uir;eis, Spemoghillrs annotus) are hibemarme, and unplate whe~~b~liry to?redation incrr~ses wirh snow acc.mdanon T~LE 21.2 Average &stances of coyote disperrai movements from established home ranges!3skoi?;cc '&!Is!986; Grse,ad Grathe 13'11: Car 5: d. ;?o6ai T, concast S'rivlk e: ~i. (i999:) reponed i change :n coyore ic:~nty ieveis dwag.war&at i.z not.le?e-dent on a zhange :n prey base. They.heonzed jiar jic rejuced ac5nr:i!eveis heipea reduce :new expendimes ;r XI area ha linured food base. Coyotes m pack 1esr more and mt-ei less eduriig h e r months tban do coyotes iivlng as matedpxs or acne ibeko3ad Wells 1981, 1386. but see Gese cr al. lp96a). Large: :ne:-i savings have been shown far i prepant fernale living m 3 ;lack &an for a iemale living aniy wth her mate (Bekoff and Wells i981). Howeve:, it is unknown mhefier Femdes livmg lo pack irc eaer ahrepoduchvely because ili!he mrrp?.raved dunng Drewaccy..-. Cayores, sinuiar to wolves, are ;vpicaily rerrionai, md?rimmiiy rernaio mrhin thrr :e.nito~. However. they also aay makc ecxmrzmioriai movezcnts into nei&bnormg teaor.es. Dispersal from the nard site mliy be into a iacant or occupied temrop In m adjacent area or they may disperse lone distanczs (Xarrison er d. 19911. In general, it is the pups ar subordinarc yearlings that typipicaily disperse (Knowiton 1972: Ne!lis and Keirh 1976; Berg and L3esness 1978: Gese et ai. 1989a. 1996a). Dispcrsal appears to be vaiuntaq a social mu nu*.- tional pressures inrensfi duing rhe winter md avauability and access to food becomes more lhired (Gese et 31. 19963; Bekoff2001a). There xeno consistent sexdiiiereuczs m dispersal dismcz(gese e:d. 19893; Han;son sr al. 199 I), alr!rough in one study, female pups moved farther than mde pups (Xellis and Keith 1976). D~spersai by juvcmles usually occurs during aurumn and early winter. rhougb same individuals do nor disperse durx~g the ijrsi yesr ma remain to prov~de cae for fitwe siblings (Kat:er 1995; Gcse et a]. 19R6a). D~spersal hc~renlon appears ro be random, and pups w~ll move %&I60 km (Bowen 1982; Gese er a!. l98ya), with record dispersai or54 km (Carbyn md Paquer 1986). Berg md CC:iesness (1978j reported mean dispersil disrmces of 48 h, w&ich occurred ar a mean rate of dour I1!m'week. Furays or ex- plor~rory rnuvmenrs xiso may occur beibre dispersal (Harrison st ai. 109 I). Increased morailty may be associated wirh &sped 3s h a i s move into imimiliai xes and low-jecunty hab~tats (Tnikowsh 1980; Pyrah 1981; Bekoffacd Weils 1986; Gese et XI. 198Ya). Observations of coyote dispersal are summarked UI Table 21.3. Home Rnnpe - and Territo~. Home ranee - size oi coyotes has been smuied throu&our their nnge. Home range size varies geagapbically ar.d sesonally (Gipson and Scaiander 1972; Be.bK md Wells 1980; Laundre and Kellrr 198-l) and within a population (Sp~ge: 1977; Bdwen 1982: Gese rt d. 1988a) (Table 22.4). vkiation in home range - size Dong resident coyotes wlrhin a popuiation depenh on energetic requirements. season, sex, physiogaphic makeup, habitat, and food disebuuon (!!muore and Kdler 1984; Gese er d. 1988ai. Home range rize aiso is inj3uenccd by social organizafion. Tiuslent individuals Addts I h ) Juverulcs Orm) Sourcc Wes Femvles Males Fcmaleb Both Scxcs Rooinson and Cummlngs 1951 Young 1951 Young!951 Robinson md Gmd 1958 bwthome 1971 Glpson and Sdander 1972 Nehs ma i(slrh 1976.hdrews md Baggess 1978 Bowen 1982 Grse 2: 21. 1989a hson :99? '?vide pgs. 'Fernaic pups. 'Exciudes &ra 51 one!c.mle hat moved a record dismce of 323.2 ion
T-LE 22 A Mean coyore home range sizes from some representanve mdes Aouiti ilrm'j Juverules and Pups ihm:) Mdes Females Maics Females Stare/Prowct Glpson and Scaiander 1972 Chesness and Bremickc: 1974.*sas.MmOmora Mlnnesora Utah Okiahoma Aibem Man- Texas Gese zr d. 19888 Windberg and Knowitan 1988,Atbnsan and Shackleton 1991 Person and Hxth 1991 Texas Bribsh Columbia Vernon: "Summer bwinter 'Brndks. dgenanon. 'Pup rearing. '~$s~ersai. SFaU. h SP~O~. range over large areas, whereas residents occupy distinct temrocies (Bowen 1982; Bekoffand Wells 1986: Gese a al. 1988a). Covotes iiving in packs who defmdunylatc c-on durvig the w&er hive much smaller, compressed home ranges than coyotes living in paus or alone (Bekoff and Wells 1980). Coyotes actively defend well-dehed territonai boundaries using direct confrontation and lndirecr means involving scent rnarhg and howhg (Camenzhd 1978; Bekoff and Wells 1980, 1986; mils andbekoff 1981; Gese and Ruff 1997, 1998; Gese 2001). T~ically, only pack members maintain and defend temrories; solimy indinduals do not (Bekoff and Weh 1980; Gese ;ma Ruff 1997,1998; Gese 2001). Fidelity to the home range area is very'high mong resident an~mais, may persist for many yeas (Kitchen e: ai. 2000b), and may be passed to successive generations. S~ in temroriai boundaries may occur in response to loss of one or both ofthe alpha pair (Camemind 1978; Gae 1998). (1 I%), rodents (9%), songbirds (S%), cattle (7%), labbits (7%), deer (5%), woodchucks (4%), goats (4%); and watermelon (4%). Korschgen (1973), in an analysis of coyote feeding habirs in Missouri, also found seasonal differences. For example, rabbits were found in 57% of coyote stomach in winterversus 14.8% in spring. Canion was foundin 15.6% of stomachs in winter versus 37.04'0 in spring. Livestock and wild ungulates often may be found in coyote stomachs and sscars as camon (Muie 1935, 1951; Ozoga andharger 1966; Korschgen 1973; Weavx Coyotes are opportunistic3 generaiist predators and ezt a variety of food item in relabon to chanses in availabiiity (Van Vwen and Thompson 1982; Todd and Ksirh 1983; bdeit e: al. 1987; Windberg and Mitchell 1990). Tney will consume food items rangng in size uon Ou11 and insects :a iar,or: ungulates and li-vesrock (Fig, 22.5). MacCracken and Hansen (1987) su:.,oested that coyotes may seizct prcy as?redicted by oprimai fbraging nodeis, but see Bcutin md CluE ii389) and MacCmcken :' '89) for review and comme3b. Gipson (1971:oud rei.ior;ll md sezonai dieerenczs in fee5nin. habm oicoyores jiving in.khsas.'he most ;omon iood item from 168 coyore sromazhs were poui~ (34561, persmons (23"%), inseas F!cLw 11.3. Coyore feeding on elk calf carcass, YeUowstone Natianal Park R-yormng. SOLICE: Phom by E. Gese.
:9--: 3kod I~.J \Ve!:s :Y8G), 'kt 3c:ui?reiz~!on an!qe,xp:i:itss. 50th jatxe ir.d tomesnc, ices oczx :Gie: lgh3: Ilisi! :%S-: icss s.6 Crorl?e ' 905). In '*-xiomnn, the 3erccXue- :i;svate src;-,~cbs ccn- 1978). C&on avruiabiliy dw.::g ~vmrcrpia~r a hge roie n coyote {oraeng md populai!on ecoicp :n norhcn rzxluns (Weaver :i):?: Todd 1'85; Gcse s:al. 19oh;i. 199hb). X;~roduiuc:ivs smtusn~y?v~s~ dueme.. :;.ore fez5ng habirs. C~ycr~s :hat ire provis;cning>r;;ls may ivxrcn :o Ix~er. more energecally '?ruduhic'?re? irems is :omcared :o son- :eproduc:~ve coyotes (Ti'd '(;owlton! 983; Hsrzscn ud Barison 13%; Sromiey 2000). Coycres in suburban 2-sa xe s~xdly adept ~t zxpioiring iuman-made food resotzcrs and w~i! ::2li$ consume dcg food?r other human-related Item (Cliargo :985. Atkmson and st.,.~,c.on.,.: 8 + :99!; h1cc:me dt ai. 1995). As a cave-r foor-'codhabir w.aiys;s; whenever 4C3I maiysis data re dsed io derelnule feeding habits, thc.nherenr orobiem of.. jrev digesnbilim - and cecovew uiood terns n:clation to zcmi prey consmfnon cm confound:cs;ii?i mi should b* considsrc? (tvexver md IiorEma i979: Kelly uld Ganon 19971. BEHAVIOR Direcr abseriation ofcovote behavior :n dx,wid is Jiffiait because oi the slusive nature ofyhr species (Zsunan mu Bmdy 1975; Bskod ZOOla). Howeve:. observarlocs of ;ovores in national varks iave orovided insight into many aspecs of :he bei:aviord ecoloa oithe species (Carncmd ;978; Bekoii' and Wdls 1980. 1081, 1986; Wells and Eekoff :981; Gcss et al.!9y6a, 19960, iyy6c; Gcse s1d Rufi- 1997. 1998: Ilcn et al. 1999). Dam on behav~orai deveiopmenr xe presented inbekod(ly72, 1974, i97%, 1978) ma BekodmdDugar!an (3000). Cavarrs show early deveiopmcnt of apsessive -- behawor cornoared to,,voi.ies dndmcst domeshc dogs; they w?ll zn-azr in sexuus 6ghrs when thsy ire only 19-24 Jays old (Knigh; :974; Bekodcr ai. 198;; BekoE mddugah7000).theeady deveiopmenr~ii~rcia~ons~ps within litters aopcm.. co iatirp 104.2 months lic~iehr - IYlSi, bur the dominance iurrvchy later m life may nor reflect the hienrchy obsrmd w i ~ thr. n inrr (Beko81977b; Knighr 1978). General behaviod panex (postures, gcsrures. ail novemmrs. facia! ~xprrssions, vocalkst~uns) uire described md discussed in K!s~man (ly66j, Fox (1970). St.kofl(iY72, 1974, 1978) Lx&t (1978). LcImrr (!37Ya, i5!7sb), and Weils and Brkofl(1981). Gait, srmce, ex and tail pasiuon. and rerracnon of 'be!ips to expose!he tecth are all ver; Importar in sacill iommunica-!,on and inay vary dependently or tc_rer!:cr, depending on hkv.dua! 'mood." Ccmparahve reviews are m :<emu. md Eisenberg (1973) -ind Kickan and Bnay (1955). Sodal Orgsnizatiuo. GcnrzCy, coyotes ire ionside::d less socral than wolves (bur see Gcse e: d. i?"f6a, l99hb). In Ycllowstone Nanonai Park hch prey biomass, :%zb survival r~tes. and lack ui 3e:secur:on packs. is the donxl;mt idult hcterose.~l pair. often reikrred :o as 'be "alpw pax (Bowen 1975; B<koffmd We% 1980, 1R86; Gese er xi. 1996,~)..\ssociate mimais w-iil remain in the?ack dnd possibly inhent or displace a member of he Sreeding pau and br-ome an d~ua theselves (Gcse cd. i?%aa). These msocates (or 3etxanunsisl paruclpare in :ctonal maintex~nc; md puo irmy (Harier 1995), bur out to rhr sxrenr of the alpha pau (Gese md RJE 11)97. ;99F: C.cse 3001). Other coyotes exist on :he landscape durside of *r rts~dent pncb ar.d are cnnsiderzd misnr or nomadic hci;.?dwds (Cme3zmd 1973; Bowe:: 1982: Gcse c d. 19SSa). Tlesi: in-~lents usmily me! ~ione over 3 large area. io ::or breed md di> not maintain a ;err:ori (BekaRand \Veils 1986; Grse and &IS l??7, 1908; Gcse 2301). Cayore pi.!- rescmbit woi<lacks!sowen IC-'3; EcSo%mtl.YCils :950: Gese -1 3i.!O?Gb,) md :r ippe%-i :hat Ji3tr-sczs ietwezn ;:qc;tcs 1-d,.~ciws =re?~atitnnvr nthe: ;ha quiirar~ve.,.!us; B.*v?h LI 'other S ~~CLS~??c:e :;ti~c!~, :hex,> com;de:- ~"le.iar,abiiiry.k observed sccrai oi:?vm~non (Li~wea 1YX; Bckod irl '.!:?:Is 1980. is$:, 1 C4h). 'a m w ires. soiirar:, :;diwduals sr: k:r;c-::!v cbic7e2,3::g md C'aesxss :?7S1 2utsiie 3ihe iesdine 1?&31. :n 3th;: ariq jr:i 1s Jachcn?.>is, Wyom:!,Canewad."- Y Y:E3esaffand'A~ci:s :?SO. 198:). md!asper,,%bem(bowez:9;8); zrcups ofcoyores a: :~moniy obse~ied.?-ey slze maybe mpom.r in i5c:lr.g :nyo:e iocidi~ -/Isowen!YS: SekoRmd We& 1980). in,-i:puiariocs -.\-here the Tqcr ire:, rtcms rhrou&our -be year are s-1 rodenrr, coycrssrs~d io he inpain orxos (3rkarimd We" i9y6: Grse e: 11. 1?8?aa). In?n"ularions vhere lxce amds are mailzbie (a.2.. ek,.. Leer; s:tns: s live mdiv:duals or as wroo. :oyores fom large gcups (Ijekoffrnd Wkas 19%: Gese er ~i. li9ba. 1995'0). However. coyore groiics.:; not nec:sssrjy fomed to capturz large prey (Gcsc s: 31. IUYSh!, :bough coyotcs oc;as~onaily nunt s 1 pair or s goup jf?h md Schwc~ner 1979; Bour/er 1957: Gcse md Grothe 1995). bsread iooperxlve goup defense xppcxs to be the mqor seiecsve force favor- :ng mc'easei soc:aii?/ (Brzer 192: Bowen 1978; Lamprec51 1978; B;kotTar@ SVr:;s 1980). However, :yen with a resident pack, dominancs plays a!ai:e role in access to fezding on czrcxses (Gese et 31. l"601 and niiuencrs thc abiiiy oi associates 10 remaln in the pack (Gesc c: si. 1996a). Imponantly, coyotes rend to be more sacid during winrc:, when ;Jrnon is a "en; importmt food resow2 ibowen 1978; Cxr..e3zind 1979; Beioff and We!ls :980). In areas without larse ;in-.. gxlzr: csrr!un,!nce~sed pack slzz aiso occurs in :he brczhiing season, which then f~iiiiures caphlrc oilxge prey (Gese e: dl. i988b). Predatory Behavior. There have been 2 kw derailed studies of ;he r;:sdatoqi behivior of -mid coyores on snail mammals (W>ils and Bekoff 1482: 3ekcfTmd W4ls 1986; Gese e! al. l996b, 1996~). P;edatory sequcnccs may be divided into a1 lear SLX cornponenu: search. ocznwriou, snk pounce, head thvclose sesrch :nro ground cover, s d rush. Pounc:ng is used mostly for capturiniing small mic:ohnc TOdcnts, whereas rhc rush is used most kequemiy on!qcr animals, such as ground squ~rreis (Bzkoffand Weils iy86: Gcse et d. l996cj..age ot' the miual, wind, habirar. and snow :ondioons (depth and hdrdnessl ail idu..3c., L coyote's ability to rice.: mu capture smallmammals (Weils mdbzkod1981; hlilrrsy mdboutm 1991; Gese i.1 I. 1996~). Coyotes generally hunt m I I mammals xione, even thoubthc yack size may be quire ixge (Gese 2: 111. 19960, 1996~). Cooperative hunting for smll mnmm~is would bc mrscient beczuse only one individual cm con- ;me ;he prey ;tern (Bowen 1981; Andelt 1985; Gese :t d. 1988b). It h~s been shown expcnmenriiy that coyotes iirpcnd on various senses to ;ocarc prey. I11 order of decreasing hponance, they are vision, audition, and oifactlon ~Xells and Lebnrr!gig), though these "pcorines.' nay change depending on envlronmenal condinons (Wells 1978'1. Pecbtirory behavic: mdhabiw ofwiidcoyotespreging on domssnc livestock have received increased ~rtention je.g., Heme 1977; Shivlk ct~l. 1996; Windberg et 31. 1997b; Xrale ctal. 1998: Sacks er al. 1999). Cznnoily 21 JI. (l97sj found :hat capnve coyotes MIsd sheep (in confnemmt) aniy in 30 ot.38 tests (5?.6:1,); defensive behavior by she:p dite~cd coyotes only 31.6?/0 of the he. Miem latency to attack was -is?! long (47 mni, with consideraoiy vanabiiiy (swndard dewauon = 18 slir.) arnun_r amc.b. Mean kiilulq rime, likewise, was cithcr long (13 :nf), and cons~dcrrlbly vanable. In mosr ;nstances, coyotes attacked,., ny biting the thros~ and the rh~q died of sudocorion. Various fac:ors influence cayore depredanan mtes on sheep, including breed of sheep, shes? mma~emrmp&ces. coyote bebavlor, snvironmenwi fx:ors. and de?re&rion management progm. In xenerd, chan~es in dnixal ilze and behavior, differences h soup cohesiveness, soclaiity, grazmy kspersion, attmnveness. and mtcmi protection ail adefeci v~inerabiity to coyare predation (sez re~nzw by Knowiton st ai. i99y). R->.dbrrx 2: A. (1997b) and Bromiey (1000a) do~mentsd that coyotes ~rxniiy 'oiled rhe mdlesr dr!is!resr lnkviduals m the dock. C~oper,u.ie buntins ai adult unpiates ii.6.. deer. dk, mrelopel by coyores me!? nas bee3 5ocllm:nred (Cahalane i9l7; L1w.e 1951; Eras : 370: Zcwyer!987\. lv?ea ;oycies mi.!; large -aid ungulates. e.lvusrmsne!?ac:~rs sue= 3s ;now de-jrb and 3ardnrss oi the zrust.. 2 2 iri~orm: :l! the IIIC~~SS iir :xure :f're mex?t igrs; ma Grothe,995 j.?:eseacr ar' he ipha?ax also appears to be impom1 in 'be
outcome oithe attack. Generally, younger anlmds do not pmicipzrs in the atiack sequent:. It appem that the mbe: oi cayores m the pack is not as imponant as which coyotes are involved m the araci. Even 2 a pack ofsevec coyotes. ady the alpha pair was hvoived in attacks on 1aree;inpuiates - - IGese andgrothe 1995'1. Also. the abuiwoi~uneulares to escape into ware: deienslve abiiities of individuals and other members of the herd or the Daren!. and numnonal scare of the individual unde: attack wiu conmbute to the outcome (Ozaga and Harger 1966; Bowye: 198:: Gese and Grorhe 1995). Communicatian. Coyotes comumcate using auditorj, visual. olfactory. and tactile cues Cehne: 1978% 1978b). Communication by howling or vocahzation is very common among coyotes (Gier 1975; Lehner 1978a. 1978b). Studies of wild coyotes have identitied many differcnr tges of vocaii7ahons (McCariey 1975: Lehner 1978% 1978b). Seasonal and die1 panems (Launcb5 1981: Walsn and Inglis 1989), lunar phase (Bender et d. 1996). and social status in the pack (Gese and R d 1998) dso influence coyote vocalization rates. Howling amang coyotes plays a role in terntonal maintenance and pack spacing. It si-4~ temtonal boundaries and the presence of alpha animals, who will confront intruders and defend the temtory (Gese and Ruff 1998). Studies on oltxctorv communication he.. scent markine) -, amone coyotes also have been conducted (Barrene and Messier 1980; Bowen andcowan 1980: Wells andbekoff 1981: Bekoff and Wells 1986: Gese and Ruff 1997; Allen st al. 1999). ~l~hacoyotes perform the majority of scent-maridng duties. Rates of scat marking vary seasonally, pack size does not lnriuence scent-marking rates, and scent marks are located more than exoected along the nenpherv of the tenitow and likelv con- -... blbute to tenitory majnrenance (Bowen and Cawan 1980; Bekoff and Wells 1986; Gese and Ruff 1997). Scent marking may be a mechamsm for sex reco@tion (BekofF 1979a) and serve as an indicator of sexual condihon, matmiry, or synchrony (Bekoff and Diamond 1976). Internal information to orient members of the resident pack (Wells and Bekoff - 19811 and alert dispersing animals of occu~ied terntones also mav be communicated via scent marks. MORTALITY ~Mortality Rates. Different-aged coyotes have difierent mortalityrares. Mortality rates depend on the level of control to which a population 1s exposed and levels of food availability (Knowlton 1972; Todd er d. 1981: rodd acdkeith 1983; Todd 1985; Gese et d. 1989a: Knowiton et d. 1999). Pups (cl year old) and yearlings (individuals 1-2 years old) tend to have the highest mortality rates (Nsllis and Keith 1976; Gese et al. 1989a). For individuals > 1 year 014 mortality rates vary geographically (Knowiton 197?; Nellis and Kzith 1976 Bowen 1978; Andrews and Boggess 1978; Gese et bi. 1989a). Mathwig (1973) found grcatest life expectancy for coyotes in Iowa at 1.5 years of age and lowest iife expec-ancy at 5.5 years. Knowlton (1972), CrabUee i1988), and Gcse er d. (1989a) reportedrelawely high sumival in 4- m 8-yearold coyotes. About 70-75% of coyote populations are 1 4 years old (see Knowlton et al. [1999! for a summary of population srudies). To maintain popdation stability, net survival of about 33-38?6 seems to be necessary (Xnowlton 1972; Nehs and Ke~tb 1976). Causes of Mortality. Most shidies indicate that human acttvity is in-.....~.~~::n~.l~g~.~r~~c~r~c~~~~~~me.lc~~~,~i~~ul~ tospirosis on coyotes is udnowo, but infected d may survive and :::vo!es >dv!sd~ 1981, I:.;: 8, ' r 2: I 1, s! i. :3h9a, Mmdbun: remain caniers for a short time. 1993, although preiationhy large cmivores (Peterson 1995; Aqo Coyotes aiso cany a Mne? of parasites. Ectoparasites com- 1998: Crabtree and Sheldon!999) and stamanon dunne crashes of monly found amang coyotes include fleas (the most common external food resources also may be substantial modty factors. b~sease also parasite), various ricks, mites, and lice: see Gier and Ameei (1959) can be a factor. especially among young or the year; pmovirus ialled and Girr et d. (1978) ibr species identiiication. Intemai parasites inseveral radio-marked pups in ~~eilowsrone igese et al. 19971. Even in clude several s~ecies oi flukes (Wematodes). tapeworms (cestodes), 5&tly explo~ted populations. mosr mortality is armburable to human intestinal worms (nematndes. ascarids), hookworms (ancylostomlds), causes. Human exploiration can be substanr.d in some coyote popuia- heartworms lfi!aroidsi. eso~haeeal worms (spinxoidsi, lungwarms nom (&owiton :9?;; &owlton et ai. 19991. Diseases and Parasites. Ssrologicd analyses for anrioodies in coyotes have shown -bt they have been eerposed to =any diseases. in \iellowstone Nanonal Pxk. prevalence of antibodies against came jmovins (CP? was 10046 for sdults, yearlings. an6 old pups (a-12 months oid), but 0% for young pups (<3 morths old) (Ges; et a!.!997 [n = 1101). In Texas. Utah, Idaho, and Colorado, >70Yb oi the coyotes bad antibodes against C?V momas et al. 1984 [n = 11841; Gese :t d. 1991 [n = 721). In Georgia, 65% of i7 coyotes had antibodies against CPV (Hoim er ai. 19925). High prevalencz of annbodies is often associated with a hi& contaeous, but nonfau! infecnon because prevalence is measured among survivors (Thomas et al. 1984). However, of 21 coyote pups implanted with radios in 1992 in Yellowsrone, 8 of I4 deaths were from CPV lnfecrion(gesee1 al. 1997). Presence of anti'~0dies against canine disremper virus was 88%, 54%'. 23%. and 096 prevalence among adults, yearlings, old pups, and young pups, respect~ei?.. in Yeliowstone National Park (Gese e: al. 1997). in Texasl Trainer and Knowlton (1968) found 37% of 33 coyotes bad antibodies to the vim; Guo e: al. (1986) reported 56% of 228. W~lliams et ai. (1988) reported that 5096 of 10 coyotes in Wyomng tesred posirive for this wus. In Geor~a, no coyotes were found to have been exposed (Holrman eta!. 1992b). he~lsnce ofcake inkcnous hepatitis virus antibodies was 97%, 82%, 54%. and 3354 for adults, yearlings; old pups, and young pups. iespecnvely. inyeuowstone Xationai Park (Grse dai. 1997). Covotes in Texas and Georpla had a lower prevalence (41-5 1 %) of virus exposure itrainer and Knowiton 1968: Holrman et al. 1992b1. The depree - to whch this virus affects coyote populations is unknown. hi YeUowstone Yauonal Park. the re valence of antibodies against the pi;:&ue bacterium Yersiniapestis was 86%. 33%, 80%. and 7% for aduits., veariines. old nuns. and voune ouns. respectivelv (Gese et d. > *..,......, 1997) Ths!ugh prevalence was similar to results in other western states (Barnes I982 in = 12.40511.., In conuast.. coyotes. in California had very low antibody prevalence (<6% of 338 coyotes sampled) (Thomas and Huehes - 19921., Covotes, mav become infected with Y Destis bv be;lng biaen by fleas or possibly by ingesting infected rodents (Tr.omas et a1 1989). Infected coyotes usually do not develop clinical signs, but develop antibody titers that last about &8 months, malang coyotes an indicator soecies for ~laeue (Bames 1982). Changes in prevalence of plague h the coyote population likely reflect prevalence of piague in the smal-mammal Drev base... low levels (45%) (~ese zt al. 1997). In Texas, Trainer and &ely Knowlton (1968) found no serological evidence of tularemia. In contrasr. 88% of the coyotes in Idaho were seropositive (Gier et al. 1978). Coyotes may conwct tularemia but are relanvely unsusceptible and will likely recover (Gier and Ameel 1959). No coyotes in Yellowstone had serologcal evidence of exposure io brucellosis, either Brucelln nbonirr or B. cnnis (Gese et al. 1997). Similarly, coyotes m Texas and Georgia had not been exposed to bruceilosis (Trainer and Knowlton 1968, Hohan et al. 1992b). Coyotes do not appear to act as si@- cant hosts for brucellosis. Prevalence of antibodies against leptosplrosis in the coyote population in Yeliowstooe was low (Gese et al. 1997). sinular to results for coyotes m Texas (Trainer and &owiton 1968) Ad Georgia (HolPnan et d. 1992b). In contrast four of nine coyotes tested in Arizona were seropositive for lepiospirosis (Drewek et d. 1981). The,&pact of lep-... and.4nce!' 1959; Hirsch and Gier 1974: kficchei and Beasom 1974; Thomron et ai. 1974: Conder and Loveless 1978; Gier er al. 1978; - -
qlzrcrr ci S. :9SS. wixson: ;: g.!':91: 2oi;n;m c: 2.!0035 I. C~yoies 3;,,, nay mp~sb!es ma ma:d ~silifer iom crrrdiovascuix- &sexes. 30:- Ju mesrjsiis it>cmtcn ;r a. I?YJ:. mange (Pe3c-n 27 21. :!?El:?PX= 'yvx~ibe:g I?OJ'.. mu cancx F.:r ie\'?ews ma :cxcncmic mdyses di s x t ~ acc ~ l memii. paaslies. see 3ekod(;97;a) and (Gier 2: ;I. 19:s). C,~:cre Jge can be tstinat;:! by cornring dmtal cemennun annuli!l;nhm ma ::=owiton 1967; Y4iis 2: al. 19'8) and coarseiy esn- ~13~26 wlrh tcorh weir (Gier!968). Before tooth sec:ionmg for cc- ;ncntli. anaiysis; 3qc class ijui-exlzs mu adults) can bc &srrn-gushed basrd on the r:!ar,ve zuip C3Vlv size usinq miiopaphy!,lknowiton and Tvhrrcrnare 2001). Roberts (1978) palnred out thar -here is variation a age de:;manon from uifferenr teem and suggesred wing canines in age ds:emuation. Eye lens weight, baniurn we:&, and herroai cnnnaction of long!endons can also be used to eshmate aee - accuntcly. Ages siyouns cuyorzs can be istimazed h m bo+ mass, body :ennrb. mu!enmh ofthr hind fool igier 1968: Bckoff and lameson. 19-5: Bmm ;r 31. 1979). The reqession equation for the body mass of hanc-rraed pups (O-30 days of age) is y = 0.1685-0.197~; for coyores 51-15.1 days of dge, the equtlon is y = 0.50-19 - 0.0449.~ (Barn- 61 al. 1979). From hand-raised coyotes, rhe resiession e p- aons for predicrmg the we:sht of!mow-ase coyotes are y = -13.57 + 50.59s (0-30 days) and y = 11.386 + 21.11.7 (31-15.1 days). Toe cond;it~ous bertieen wclzht and age for!&3o days md 3 ;-15.i days xc 0.999 md 0.995. respectively (Bamum et 11. 19-99). ECONOlVEC ST.~LTUS AYi) hi.lu.\gemx.w Coyotes arz,ncrlms of success. By takn~ ldvanrage of poorly devised dorncst~car,un orscrlces that lefi.most livestock virtualiv defenseless a~ninst ;r:~.on, coyores have esmblisbcd a reputation for eaic;ent and r12cnve zrcdation. Sheen havcbeenseicaivelv bredto~ioduc:anuluna!s that are siilted to pacticulm ';usbandy pracric~s, regons, conditions, md cuihr~l tates as well as to piov:de food and fiber (Knowiton st il. 1999) DiiTerenc.5 in group cohes~vcncss. sociality, gaxng disperslun, loc3tion; nnenrivcncss, mobiiiv, behavior, and maternal protecuon all 2Ec:cct thcx vulnerability to prcdatlon (Giuesmg et dl. i980; Blakesiey and McGrcw 198.1; Knowlton 21 ai. l999l. hi~or;antly. nor ~ll coyores :GI! sheep. Some are never exposed to shecp (Wagner 198Y), whereas others do not develop sheep-hilin~ rendenc:es jlr.s. Fish md Wildlife Sem~cr 1978; T b and Connaily IYXU). Shiv,& 6r 21. (1996), Canner er a!. (IYYX), :ad Sac:- ct 31. (1999) described radio-coilartd coyotes nesr sheep \nth k\r-!mbs brmg killel;; losses were usually ircr,butrd ;o thc brerding, tentodal cayores. In conrrasr, Connolly (1'188) md Windbcrg r: d. (1997b) E- oorrsd a hi=! numnbcr oicovorss thm killcd and consumed!ivcsrock. In the internountab West. Till and buw.ton i 1985) md Bromiey (20001 dzmonsuatrd ttut aduit coyales wirh pups were more lkeiy to hil iambs thanadults \i;thourpups. In dunnzsr. renironal ioyotes innoa-coaswl CdiLiibmia laii Imbs as soon= hrv nbccamc~vailabie :ndecember and January, oursldz the pup-rcd~g srson (Connrr 1995; Sic!ti 1996). Economics of Coyote Predation. Tne mount of :ivestock!osses that produce3 imibure :o coyotes is J sonrenrious issue. Wildlife protrcnonsdvocztes :iaim~ht ie.u or nu depredarions actuaily acc.~, where= ;rroduce~s kdicare!asses From cayates threaren their economc livclihood!caine m dl. 1973: Ge- et 31. 197'; CS. Fish md Wildlife SzMce i?7s: Wagner 1988; Knowiton er ai. i094). In iiruanons where here was prehror :onnoi. losses of sneep ro cayotes were :.il'.o~o far Imbs =,ld 0.1&2.0?/0 for ewes (US. Fish and Wiidlife Semic: 19781. h %>ward 19-6; 3rawiz:i I?--: Heunr 197 :vllmoz 1977: Llc.idoo and :ie!:e~o\v 19-8; O'i7~r e: ii. :!JP>. In :?%. :he zcanomic value ui domes::^ sheep 10s; ro pre&ron was reporrsd is $17.7.milion 11-5. Fey-z-r jf >.rri.-!rure 19q5i. ie=irr m6 S:;m&e (:978) md L~.S De3m:s::: ois=$;?liture i:?95! rnanzd -hr c3yotes wc~e rsspc, n>,o:c $1 he ;lajcr;.i :i 111 she:? lost :o preron. iiowe.ie:, ;r is hpocjr.: :o corsiccr :hat coyctz prehnon is ilot 'ie najor c3ue oi :css;s Cis. Fish ma 'Xildiiie Serilce!9-8: Bekaii 1979b). Eariy s7 1:. ( 197.1) found thzr -7% ai the rotai cc3nonic ;lass oi iherp k :3-%1?7l was Lue :a Asease (43?'0), w.spec:fisi zauses 131?6j. md pre&rcrr C3"/oi. Depredation \.lsnszement Protecting 1ives:cck or wlidife species iom :oyores :ccn;;s ;ons~de:anon ut';qal. soc:d, sconomic. biolosicd. :;c:m~wi. 5:bcal fac:ors (Stener an3 Shumke i97s; Wade 13-5; Bekoff 1908. 20Olb; fficwlton et ai. I'J991. Succrssliii resoiut:on ofcodic:~ should cansider rhe efficac:~. seiecuvlty, and ejicirncy ot' vanous proc:dures. S;.re.al te-.ki~es xe avaiiabie?or reducmg czyore depredano~s on!ives:ac!c (Siemer md Shumake 1978; Wade 1978: FA 1590; LS. Cesamncnr oi.4gnc.dlure 199.r). Limy tecnniqw's are within rhe upcranonal pur,,ie.w of the producers, such is nunie:hal tcchnlqces. ijthers invoivs lethal co~noi in either a preventative or ccrrecnve context. Procedues ;Oat are more bsmp in ebects on narurai sysrrmj ire rypiczlly prrftrrd. Fd1 11990),.4,udeit (1996). and K;icwiicn 2: al. (1999) :ev~e..vca mmy of the rechiques for reducine coyore depredacons. Y\ronremu\ai Techniques. Szved rec!mio,ucs have been used to detsr coyores %om attachnq livesrock (Linhan 1984a: Wagner 1988; Fall 1990; Green et il. 199.'). Wberex same methods have beensilccrssful, others are complete i3ililres. Knowlton a 21. (19%)) suggesied several husbmdry pncrlces tksi may reducr depredations, including combing or concennstki. - Bocks dunnv -. uefiods of vuhcrabdilv, usme - herders. shed :mbmg, removxiig lives:ock careon from games, synckom~g binhmg, and kcepmg young mais in areas with little cover andlor ncrrrrsr to hum=: icnviv (Iiobr! r: ai.!?91: Wagner 19831. Scvemi kncs.. sf 3rorcsrive fcnc:nq c:n exciude 31 reduce coyoi- use ofu~ are.>!or idesta and Cropsey 1978; Llnnart 21 al. 1982: Sheiton IYZ4, 1987; Vass 3ndThe;ldc I 9RS), but few are"coyorcproof." Fnghtcning dcvic;s!hat:nit blvsrs oflight oarsoundhave de:ened coyote Drehtion on sher~ in fenced pasrures (Llnnart er al. 1982, 1984; Linhart 198lbj and on open range ili&.art e: ai. 1992); howe-ier, covatrs often become habimared to such devices (Knowiton et a1. 1999). Use of,pard lolmais may reducz damage by coyotes. Dogs, Uamas, and donk;:,s ormulcs xe used 35!ivesrock guards jlinhm. e-l d. 1979: Coppinge: et xi. 1983; Grrrn and Woodruff 1983, 1987; Black and Grecn i984; Green ;:.II. 1984; Tjnm and Sc.htnidt 1989; Powell i993, Conner 1995). There arc no re3eue3rs md!emed aversions rhatwill deter coyare predation. A vanety ui'~:;tatory olfactory, andmrafmgproaucta ;have been tesred. Food -onsummon mav be reduczd (Hoover 19961, bur predarion is nor (Lehner 137; Bums and Lhson 1997). investigations of condinonedmste,lverslan usmg iihiwn ~uocde showednuxedresults. Some xscarcherr reported success arrcducmg food consumpnon (e.g., Gustavson st xi. 1974. 1981; Ellins md 41m 198:; Fonhmm-Quick e: 31.!985a, 1985b), whereas orhers repaired no ieduction oi'predation bv. couores. ibekoe:97i; C-nover 5 ai. 1977. Bum 1980;!983; Bums and Connolly 1980, 1985; Bourne ma Dorrmcz 1982). Sttemats to reduc; covote reproducnve rdres and popuiabon lev- eis ]lave been :nvesr;gateu with rhe assum~con that iewer Gayotes will rzsuit in fewer i;e3re&iions {Knowiton sr d. 1999). Copre repraducnon,was reducea'sing diethyistiibesreroi [Baiser 106i; Laban cr ai. 1908:. bur was :~mcmcncd withour effcc:ive bair ieiive? svstems..qternar.veiy, reoroaucrlve inrede:ence usmg jret51iz3tion 3f temto- nxi. breedin?: - ca\otesrn;lv. bem,-~edlv;~~av:o reduce de~rehtions by coyarcs ~Brodcy 2000) by ib&g ;he pre.&rov bciavtor of adults when nro\.is;orim!g young (Tii1 md ~kowiton 1983). With respect ro chernosrerianrs. Sotin rcqianon ifedcrai wd S~IS i 2nd disnburion xc prohemrn:c, SteiXug 2: 31. 1978). Cz,yote Remuvd Techniques. X%7sen ncnrzmcwi :ccimli;ues do -or stop deprehcioos, rmoving one or more soyores may acheve
- mananernenr obiec5es mowiton e: ai. 1999). es~cciaiiv ii the removed anlnais are the "problem individuals" (Linnd 5: a1 1999). Removlnn one orwo :n&vudi;als m a amall area ii.e., corrective ccmol> may stop the problem. whereas mother cases,?opuiationreaucnan may be warranted Knowlron et al. (1999) suggesteathat selection of the appropnate method shouid corsiderthe name oi&e probiem presence of a hsroricai parreni. sue of the area season oithe year, tuning, efficacy se!ecn~ry, -5c:eacy: anaatumal weifare considerations (Bexoff 1998, 200ib). On small areas where specific coyotes pose the most &mediate rik cahg and shoo~g can selectively remove coyotes killing livesrock (Coolahan 19901. bur requues conectiv identifvine. - rhe areas used by the hllers. One method thar mav be the most selecnve for remome - coyotes. responsible for depredations is the use of the livesrock protection collar fmcbnde 1974, 1982; Burns et al. 1988, 1996; Connoilv and Burns 1990; Connolly 1993; Rollins 1995). These devices reiease a toxic chemicd into a coyote's mouth when the coyote attacks and punctnres the collar on a sheep's neck. They are registered by the Envlromenral Protection Agency in seven states and require approved Wining and accountability programs (Moore 1985: Knowlton e: al. 1099). Coyotes causing depredations also canbe removed withuaps, snares, andm44 devices. but tbe selectivity for Che offending coyote(s) is lower (Brand et ai. 1993. Most depredanons are attributed to terntonal, dominant coyotes (Till andknowlton 1983; Sack 1996). Withthese tcmitorizs, coyotes are less winetable to capture devices because of thelr farmliarity of the area (Hams 1983; Windberg and Knowlton 1990: Windberg 19963, which makes removal of offending lndividuds dficult (Comer et at. 1998: Sacks et al. ip99). Aerial hunting in winter is efficient and effect~ve (Wagner and Conover 1999). However. concerns regarding the safeq of pilorr and gunners, seiect~vity of anunals, aod economtcs put see W-agner and Conover 1999) are a matte: ofpublic debate (e.g., Finkel 1999). Coyote Population Reduction. There may be sitwations in which a reduction in the number of coyotes requires serious consideration, including instances where coyotes pose a risk to other wildlife species, the spread of infectious diseases needs to be curtaiied or predation on iivestock needs to be re vented when more benim tecbniaues have been ineffective (~nowlton et ai. 1999). population reduction programs are mast effective when conducted &er the dominance and territoriai patterns are estabiished far the coming breeding period and before whelping, to prevent other breeding pairs from becommg established and producing offspring that season (Know-lton 1972: Connolly 1978; Knowlton et ai. 1999). 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Mavernmtr of breedme co?oirs with emphals on den sire &iaoonsbpa. Journal of Mamma~oG 60:568-75..h&k W 5. I. G..Gc. E F. :ecwlto% md K Cxdwei!. 198- Varmon lo cavore &e:s asauiarsdw;b icaon and ruccessionji changes m veaemnon. Journal aiwiidliie Manageaent 51:273-7:..h&wvs. R. D.. ma E. K. Bosgess. 1978. Ecaiogy ai coyotes m Iowa. rases 21965 in M. aekorf ed. Coyotes: Biology, behmar and managemcnr. Academc Press. NCW York..Qo, W. >l. 1998.?'he effecr: of recoio-g wolves oc coyote popuiaoons. movcmenrr, behaviors. and food habm Dissemnon. Umven~ry of.uontana Missoula. hrjo. W M.. and D. H. Plcsche:. 1999. Behavioral resconses ofcoyores to wolf iecaiomanon in northwesrern Mangna. Canadian J o d of Zoology ;i:i91'%27. Asdell S. A. 1964 Panems of mammaiian reproduction. Cornell Gniveriity hss. irhaca, NY Aths. D. L. I978 Evolution andmorphology oftbe coyore h. Pages 17-35 m 1. Bekoii, ed. Coyotes: Biology, behamor andmmqement Acadermc PKss, New York Atians. D. L., and L. S. Dillon. 1971. 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C. 1986. Socral ecology and bchavlor of coyotes. Advances m the Studr of Benavioi 16:251-338. Bckoff. M.. H. L. W. and J. B. ~Minon. 1975. Behavlorai manomy in mds by discruninant iuncaon analysis. Science!90:!22>25. BekoE. M.. bi. TyrreU M, V E Lipen. an6 R A. Jmieson 1981. Fighting patterns ix young coyora: hitiaaan, escalanon, and assesmem. Agggcss~ve Behior i:z2.w.
3e2ae~ 3 I.. T. >I. %I-e. lcdi 41. Zcghm. 3% 2-ccnjiiion &~:ezces ;jyarc (Im!r:nrrnrui.ii;w!ing. imcnc3nltidlanlls~~iis;.30.111-i-,., 3: W 5.. ma h A. C.csress. :Pi4 izoiug.>i ;a)orss n nor3ez iinr.inr.sam.?=ees 1:0--17 :n M. Bcrof. $3. Cayxes: Bioioe. beenwar md nr-agemmi..4wilcrmc Eess. Yew Ybrk. 3crge:. J. :O-3. '?raaior imssmmr' LS I liefensive smtsg..aenwn Mid-!and Ydlsr :Oi:!9'-39 3:nir H. I.. ma 1. 5. Green. 1081. Na>,a!a ise i;nu;c'-bred doqs for man- :qcmem aipreurarj. Joumai oiilaog? Li-qemcnt j8:i 115. ~biaizy, i. 5. ma I. C..LlcGrew 1984. Dt2s:ennni uinsracciq ~i1amor :a cayore?i;&non. ippiicd ha! Benancr Sc:mcc 1::34%5;. aaggsss. E. K.. ad%.x Henderrm. ;?X. Rcgonni rve;i.hts uibas coyares. Trriacaoos af!he hsas Academy oi Science 8O:?LdO. Bourne. i. mdll J Dorimcr.!982..A field resr oilitlxun ;Sonde avcrslon ro rccuc- ;wote?reunon Jn domesiic me=. Journal oiwildllic Imagernes 16.235-39. Bourn. 5.. ind B. D. Cluff. 1989. 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Cappinger, X.. J. Larcnz, aid L. Cdppinge:. 1988'. huoducmg iivestuck gm~dmc di-s io sncrp and joar productn. Procecsingr oi the Eaaem Wlidliie Damage Canuoi Canference 1 :!29-32. Cmbtrer. 'i L. i48y Soc:ademogapiiy of an uncxplo~~cd ioyute populzian. Dnsenatian, UrilversiN of Idaho. 4lurcow Cisbur-, R. I., md i. W Sheldon. IP9R Cayotvotes md cad Zaeristcncc m Yc!loiustonc. Pngcs 127-j3 in T. W Clark A. P Curie:. S. C. Llmti, md F. hr:m. 5"s. Cmivores n ecosystems. Ydc L~nlvcrstp Press, New Haven. CT. Crooks, K. R.. and M. E. Sod= 1999. Mesopredator reirasc mi avliaunal txtmcnous ma hgmsnred aysrem. Nature 400:563-66. Cmmghm, V 3.. andr D. Dm'or*. 1970. Rcccnr coyorencord iromflonh Qu;irre;iy Jbunai dfhc Florida.<cadeny of Sc~c-cr ;3:279-30. C:?hcr, B. 1.. and K.. Sprnccr. 1993. Compc!i6-ie inrerjctions benuem coverer and San Joaqum kt foxes. Jo-d aimmaiog 79:2OL211. Davisou. R. P 1980. The clikt of cxda~ration an some pardmcrerr oi coyote papuinnoris i)~rsmarion. Utnh Shte Umveniy, Logan. Dc C;ilcst~ D. 5.. M. G. Cropsey 1978. Fiela test oia eoyoie~prooifence. Wadlife Sucieq Bullern 6:25&59. DeLorenzo, D. G., md V W. Huward, ir 1976. ivaluanao of sheep Lasses on a rang lmbmg openoan wirhour predarar canzai.n sourhrssrern New Mex;co. Find rtponza :he L.S. Fish ma Wild~fe Scmicc. Denver Widlife Rcscarch Cente; New Mexlco State Umveniq, Laa Crwes. D~ce, L. R. 1942. A imly of dus<uyore hybnds. Journal of Mamalog 23:186-92. Drcwek. 1, ir, T. H. Uoun; K. J. 'finurnan. and E. i. Bic.helL 1981 Scrolodc svldencr ai1eprospaosis m a routh~m;\nzon~ coyore popuiat~on. Jomd ai'htidlife Disessrs 17:33-37. Cads, h 3.!948 Ecropmsltes iiam a scnrts uftexas coyores. J o d ofmmmalag 29:26&7i Cxly, I. 0.. J. C. Raethcii, md 0. R. Brewer 1974..h ccanarmc study of precnon h the!dho m.gc sheep mdus~, 1970-71 pioducjun qcle ipraeress Rsoon No. 182:. Idmo hunr;itud - Rescmcn. Um~enlri oi idah;, Moscow E!&r, 'N R., mnc.ll. iiqdm. 1977. Us;of&sc~mmrhuncuonmraxonomc ie:;.manon oi cmds tom Missow.. Journal ui Mummaiac 2!:17- ilii. 41. W! 990. Canno1 oi ~oyatc depledation an livestock?regress in resemh md dcr,eiopmenr. Proceedings of rhc Venebmre nest Cooference 143-51. S;&n!, M. 1999. TIC oicmare survivor Audooan IOl:5Z-S9. Fisher. J. i9is The pialns joy moves cast Nanooni 'Niidbe 13!Li6. Fonhman-Quck 3. L., C. R. Gusravsan. md K. W Xusitmk 1985a. Coyote isnuol and msie &yesion. ppenrc 6:2534a. Fonbmm-Quck D. L.. C. R Guswvsan. and Y W. Rusmak. 1985. Cayores md 135:~ 3ven10n: 73~ m~bors'.qly Appr~e?:lSL30. FOX. >L. V!Q7O A iar.~ar~rive ~:u(iy aia; ~sveioprr.mm ai5c:al ;xpressions m zamdr: 'Yoii. ;ayore. foxes. Br5~iv1or ;6.19--3. i-ale:. :. K. md L. 3. Kt:&. 198:. Non-.>\en~npns mues 2i cayores md solves in norrhmstern.ube.m. Jomd aimammaiog 62:403-5.
Hoors:. 3 :??6 5i;:nenrss ai-iainni; roezmal mmnrr ar redusag egg ~;n?unipnoa iy xmalim pemron: in :xpermencd maiysis. *Icsa,. ~trn - Si;is Univernv, hgm. Zanl. j 'T.. ma P V L:hs:. :^5. Si~ioplc irnani~rj in the coyare (Gnu Icinrm:. loesi dfic$mpm~ve Physirriogy md Psycnotag 39::070-:6. %ouston. 3. 3. 19-3. Fk IS.Inre:-spch:i ioad kr -mores m norrhern k!- :owsroiie Ndnona Pinr. ;omd oi-lppiiei Ecoiog 1565341. Gowars \V. E. 1944 4 mrms :o iisnngusn jrdis ai zcyores md 2omes.c mu,- Jcmzl oil1unm;uo~ 50::03-7:. lacdon. 3. H. T 195:.?n3. C:assuicztir,on ofrhe scrs dicayores. Pqrs 237-341 :iz 5. Young mu 3. H. T. Jackson. cds. X,r cieve: sayare. 'k~&fe lanqcncac!mnsiimze, INashmgron. DC. :(;!y. 3. T, md E. 0. imn. :99:. 2fec:s of prey size, meal rlzc, mesi cornposioan md dady fiequenq ai <ding dn the recovrzy oi mdenr rernsm iron camvare sia;s. Cainndzan Iounai oizaoiog 75:lYIl-i- Kcnneiiv.,. J.!. :97Z Cworc ;-.crcducnon Pu; 1. il;e dumnon si.he seernarcgenic c:ic!e and ;?~did:.::~d spm mspon. Journal oixzproducave Fer"liy 3 1:!65-7C... ~,nnc:ly,j. J. 19-8 Cayore:cumdocnon.Pzges 73-92 in11 Bckod rd Cqores: Bluicay, bc%;inor md mmagemsni. sademc Press, Xew "_ark K~mcUy, J, I, a d I. D. Robsns, i909, Femlitv oi;ayore-ioe,. hybnds...:oumd oii~awn;iiog jo:a;o-;1..,..utchcn. -4. >I.. 5..M. Ccsc. 2nd E. R. Schauicr 1999 Resowiz oamnanine bcr.vccn cliu-o:cs md *wit foxes: Spacc, me. md dic:. Cmadian Jounvl oizooio~ 77: ih4j-56. Kirchm, A. M., E. M. Grsc. md E. il Schsustc:. 3M)Oa. Cihm~es m coyote ~cnviq p:ltrm due :o :enucrd cxposus :o ;human prrseciiimn. Cminadim Jaurnai of Zuolou 78.355-37. :Ctch<n..\. >I., E. LL. Gerr, and E. K. Schauster 2000b Long-rerm ipaiial stabliiy oicayore ( C~U!oron?) home rmgrs ir. iouthe%rcm C~ioraao. Canadla Jawnal oizaoiugy 78:45844. Zcirn.m. D. i2 1966 Swnt rnivikng in Cmdze. Symposium ui;hr Zao~ogcni Suciery of London 18:167-77. Klcman. D G., and C. A. Brsdy 197s Cnyorz behavior in the conrzxr oifrecrm caninlci:ssarch: Problems and per. icc:ivcs. Pages i63-88 :n LL. aekoif ;d. C~yotcs. araio::: Klem. D. ti.. ma!. behawor mdmnagcneni.ic~dcierolc Press. New Yak. F. Elsmbcrn. 1973. Carnoanson of cad ind felid socriii Utah Stare UGvers~ry, Logan. :(nowiton. i; E. i972. Prellmi- srerprctations oiioyore papulanon mcchm- ~cs ~5th some xmagmcnr mpiic~ouns. ;ourmi uf Wildlire bima~ement 36369-82..Yl~awltoa, F. F., and E..LL. Gcse. iy95. Cayate papuiaaon processes re.risired Pages I 4 in D. Rollins. C. Richarcson, 1 Bldenshp, K. Caon. md S. Hake, sds. Cayores ;n illc suarhwerr: -4 compendium of our hiowledge. Texas Parks md Wiidiie DepanmenL Austin. Rnowkon. F F, mdl. C. Stauitvr. 1983. Coyote populanonmrchsolcs: lncther look. hgcs 93-1 11 in X. Bmcll, 3. 5. Lsismun. md 1 LI.?;rk cds. Symposium on n ~ i re;nliui~on d aiwiidll5 pupuim>ons. Foissr, Wudiie, md Rmsc Ex~cnmcnr Stahon. Umverritv oilhho. >Loscow. 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I93 Tie jlii UI~JS:C~ iily~ii-s ICau ;umon3):,;ommea~...oumi af!dmzaaiog~ :A.:ll72--J... L~umlrr. J. tv I98 :. Tcniord.r.ir.;inon m isyoie iccdizadon ntes. : o d oi Wddhie.memen<-15:767+9. L~unarc. '.'V., ma 3 L. K?Usr 198; Some-imge <lze af'zoyoies:.a cxniai re..?cx. loma, ~iwiililifr Mmageaex 18:!17-39. Lawreact. 3. ma 'F 3. 3osscn. 1967 Ltultipie chmsrer mdysia af C~nis.'upiir hmns and,<amilinrs. wch 1 discdsnon ioi me re:atiawiups ai C~nw nyer Amencan iooiog?s;-:133-~1. Lai~eacc. 3. md 'N E. 3osser. :964 Tllr craniai n-idencz afhybndu3aon nnew Engiana C~nir. Biw~crs 350:l-13. Lawreace. 3.. md'v 3. F3assen. 1975. Rcist~onshps oinilnh Amencan Canrr 3hom-a by imuiupie ciaractervlalysis ci~elemedpopulaoocs.?lges -3-36.n \.I. W Fox, ed 3 c,mid imds. Vm Yarmd Rcinhold Xcw Yom. L e b a Y., A. Eisehwcr 8. Sanscn. L. D. 41r;i. R. 0.?;rrrran, P ; P. Gum. mdr :C Wdyc :YYi.inuogess;on~lfcoyorernrorhondndD*iA Into jyrnpalmc.hcricm gray wol~opaianons. 3,oiunon 45:lW :9. Lzhe:. 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Bmmm, md D. 5. 3dsc:. 1968 Fieid c-ialuation of m anufemlrj agem. sulberterol. for hblung coyote qruduction. Tiansac - tions of rhc 4onh.hencan WiiGic Canirrencz 33:316-76. Lnhan, S. B.. R. T. Sterner, T. C. Cm.gm, and D. R. Henne. 1979. Roman- doi guard dogs reduce shcep losses Id C3yOicS:.l prel-ary cvalmtion. Joumni oirmse Management 3223841 T.mnan, S. B.. J 3. Robcns. md ti.!. Dascn. 1982. Elecrric fencmg reducss coyorrprcdilnon dn paswedsheep. lorn1 ofhgc 4lmgemcm 25276-91 Linhan S. B., R. i Sterner, G. I Dasch, and J. W Thade. 1984. EBicncy of lidr and sound stimuli %ircducmg coyorr ;rreda!ian upon pasmd sheep. Prorecr~on Ecology b:is-84. Linhan. S. B., G.!. Dasch. R. R.!ohnson, J. D. Robens, andc. JP~ckham. 1942~ Elecrromc.h&rhrcliing devices ior reducmg coyote depredation oo domestic she:?: EEC~LY under rmgc conditions md operationd use. Practrdingr oithe Vertebnce Pesr Canicicnce 15:386-92. Lmell, J. D. C., 1. Odden, LL. E. Smrh R. mrs, and I E. Swenson. i999. 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