Bekoff y Gese 2003 Coyote (Canis latrans)

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/APHIS/WS National Wildlife Research Center, eric.gese@usu.edu
Follow this and additional works at: https://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.
https://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~,M m q a c n t and Cm-Uoq
2d d O o n Edrred by G. A Feidhamer B
C. Thompson and J 4 Chapman. Johns Hopkrns Ihivenir). Press. Bakimure, Llqlund
Coyote
Marc Bekoff
Eric M.Gese
Cunls latram
NOMENCLATURE
~ o m o SAKES.
v
Coyote, brush wolf, prairie waif, 9e.d woif, Steppen&f. iobu, . h e n c m jackal; the word coyote means 'barking dog" and
,,&en
tiom the .\rrec word coyor1
~ c m m i Nhm.
c
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
swldespreadin S a n h .America. The Eastem coyote, formerly reimed to
1s me3 e w Enziand cmld appears :o be a recent i m m i g a t , 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. L e h a n s: al. (1991) and W h ~ md
e Lzhman (1992) presented
cvidence that coyotes have interbred with wolves in arms ,where wolves
arc m e m a conspec5c mates may not be re~dilyava~lable.
Subspecies. There u e 19 recognized subspecies of C. liltrum. Howzwr. because of the mob~liwof covores, the mteqnty of individual
scbspecies and :heir m o n o t i c uhh& are quest~on;ible(liowak1978).
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).
Coyotes are Nearctic canids. They occupy many dive:se habitars between about IO"N latitude (Costa Rica) and 70"N latitude (norrhem
.Uaska). X ~ e yare found h u b o u t 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 h u m m (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 x e 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 c x
Uually be afierentiated (although overla? andhybddization can u c m :
' d ~ ~ ls ge r o l o ~ cpume:ers, ienral ~.i.xac:erisncs. c m a l measuremezts. ne-oanaromcai ie~nues.diame:er of the nose ? a d diame:er
'frhe 'hind :oat ?ad, :ar !engh, m c k size. 3?r.ue irnph. ?elqe. behav:or. m o :ezrccs ;for re\ie.xs. see L~wrenc;md Bossen 1967: Bekoff
19- 3 % . Eider
. and &yarn 1977>.For exampie, coyotes x e typically
'
F r c m 22.1. Dismbution o i h r coyote !,Cnnlsiofram).
smaller than gray wolves (Table 22.11, thus the nose pad (about 25 nun
in diameter) and hind foot pads (less than 32 miare cor;espondiigiy
smaller.
Coyotes may be differentiated from dogs w i g the ratio ofpalatal
wdtb (distance beween the h e r mar& of the dveoii ofthe upper
iirst molarsi to rhe leu& ofthe uppermolar toothrow (from the anterior
m x g n 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
p a l a d 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 , w s , 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
~ i s ohave .:=ore pronounczd sadrtal crest -h
coyotes. ?iIuitivanate
;e:iruques : w e cleariy shorn
coyotes, wolves, and dogs can be
dizerennated anaromcally !Lawrence and Bosserr 1967; Eider m d
Bayden 197;) and behawiomlly (Bekoffer al. 1975; BekoE1978) and
467
TABLE22.: Representanve mean coyote we~ghtskom a vanety of locales
Juveniles Cxg:
Add* &g)
Sowe
,Males
Females
Xaier
Eeades
Gier 1968
Hamhame 1971
Richens and H q e 1974
Andrean and Boggess 1978
Berg and Chesness 1978
Boggess and Henderson 1978
B m n 1978
L ~ G t i s1978
, U m y md Bouim 1991
Thurber and Peterson 1991
Windberg et al. 1991
Podle et al. 1995
Windberg 1995
14.1
11.2
15.8
13.4
12-13
13.1
13.1
14.7
10.3
12.9
10.6-11.4
12.5-16.0
10.9-11.0
11.8
9.8
13.7
11.4
11-I?
11.0
11.5
12.1
8.0
11.1
9.1-9.6
11.&14.2
-
-
-
provide for more rigorous analyses than do univariate methods. Further
rehement of genetic t e c h s u e s 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
size varies with geographic lockle &d subspecies
about 400 --long.
(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 ( S c h e
and Lavlgne 1987, ~ h u r b e r a n dPeterson 1991), but see Lanwkre
and CrEte (1993) and Peterson and Thurber (1993) for rev~ewsand
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 ~ l and
e 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 9 i c a l l y 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
-
l&ll
-
10
-
Sf~*Wince
Kansas
M Cmomja
Mame
Iowa
Mhmesara
Kaasas
-
-
10.&10.8
8.G8.9
Yukoo
Alaska
Te~as
8.0-8.4
Texas
-
8.8-9.2
-
Aikm
Oklahoma
Quebec
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 kFizPetersen
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 r o k o n e b ~ & s(Horn and L e h e i
1975).
REPRODUCTION
Genetics and Hybridization. The coyotehas 38 paus ofchromosomes
(Wurster and Benkchke 1968). The autosomes are acrocenmc or teiocenmc and the sex chmmos&mesare 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; M e y 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 a r e p r o
ductive hormones and reproductive behaviors in coyotes have been dereproductive 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 = , 2 1 8 , >.05).
~
Kennelly (1971, 1978) documented spermatogenesis and the estrous qcie. Proestrus lasrs about 2-3 months and e m up to 1 0 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
(Grandage 1972). Juvenile
penis is locked in the female's vamales and fenales are able to breed (see below), although juvenile
females may d a t e less than adult females.
.J..],1 L~:-.,..,re naunr:Ag itrsrr.co by X r mde ;nu ~ : i-la2
l her-ali :c one
j:dc &%en %~n.g. ;he male rreps ovsr the k r m i e ' s 5acx an11&e sauple
reaalr. lccksd i t :SO' for i p io 25 7112. i h i same ?axn a y i r e d i o m
ye= :o year, bur nor neitssaciy for !Ze. ;\iui! csyores may nmw
p a boc& and \wheip Jr slre pups even wnen !&I2 y e m oiage iGese
i990).
Pregnancy Rare. Cestntion. and Litter Size. Tbe ?er;:suge of ;emales dia b r e d m a gvex :,exu.v&<eswt;h :ocai cmdidons (Gier i 968;
FL?owlton 19-2; Gipsan ?: i.'9775: Gese 6r xi. !989a; K?owlton m d
Gese 1995). Fuod slipply :s usudly :he prime izi:or; in :ood yeus.
nore females, especially j s x k g s . breed (Cie: '.968; Kilowiton m a
Gssc :995). Uslliily, about 6&90!6 o i a d d t :emales m d &:09'i or' female year!ings .~:llproduce litters @howlton 1972; Gese et a]. 1989a;
.Knowi;on st 21. 1949). Tne gestest m u a l 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
m d 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 ~ E i c t sof h o d on litter size. Duing ; ~ e x so i h g h rodent
dtnsir): mem liner size is hi&er (S.84.2: than h years u i 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 i t a]. 1981; Todd and Keith 1933; C3Donoghuee: l.1997). Gcs;
er al. (1996a) hund an increxse h coyote lice: sue afier coid snowy
winters ic Yellowstone Yanonai Park. Wiiren w t b harsh temperatures
and lie? snow incre3srd !he number of mgu1;lte carcasses (because of
soy016 predation and ~ ~ e a t h eavailable
r)
to o d a t m g females. The sex
ratio oflittcrs is generally i:l, though it may be skewed toward females
in a r e s oibigh ~xploiration&owlton 1972;Berg and Chcsness 1973;
Keiman and Brady 1978; Gese et d. 1989ai.
12.3. S M I of the coyote (Cmms lirianr). From tap ro bunom: laced
eew o i c r m u m i~rcnlmew oimanciiblc. hnd view a i c m u m vclltni
vie;" o i c m m . dorsal n e w aimandr'nle.
F:c-
blaies and females show annual q ~ c i i cchanges m rqroductive
anatomy =d phys:ology (Kennelly 1978). Females u e seasonally monestrus, showing one period of ' % ~ J I "per year, u s d l y dunng January
and blxch. depending on ~eogmphicl o c ~ l e(Hamien 1939: Gier 1968;
Kcnneily 1978).
P3iT Bonding. Tae dynamics of h e t e ~ o s e x dpair bonding are not
known ior d d cayores in any detal. Data on cqtive indinduals do
not a p p r u to &Err sipficantly Erom those forw4dcogores. Caunship
nay b e ~ a
ns 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 b e r m m g o f ~ . cbreedins r m s m iBekotYau31amona :9-6: Wsils
m d a&d 198:: E C C ~ ~ S!ZOO:GCSCm a RUI? 1%- :%x). ~ u r = y
exiv. surirs
. 31 comsaip. -he maie ';cc3mes lnc1~3~in'~i:i
~mc:ed :o
h e i'cmje's m e and feczs. R J c n h e femaie is re3dy ro copulate, she
;o :he base ofthe tii1 is 2bour 160 mm (Gier 1968: ~ekofT&d~ a m i e s a n
1975).Eyes opeontabout 14 days. Tetherupton thesverxyc as follows:
upper c u r s , day 14; lower c ~ x n e sind upper incison, day 15: m d
lower incisors, day 16 (Bekarl and J m e s o n 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,' u s d l y siblings fram s prcxous y e x (Brkoorf and
Wells 1980: Hatier 1995). Food ~rovisioningofthe female may occm
dunng ihe nursmg period ( ~ a r k r1995). 6 c l p h a male m d orbrr
ass~ciateswill dso help to r e x the young by providing food a s t h e
young g o w older (Hatier 1995). Young are ~ e a n e at
d about 5-7 wee.h
(Snow 1967). Tney b e ~ mto sat solid food at about 3 weeks. when
the caregvers reprgmte semisolid ibod. Beween b m h and week 8,
averaxe weight increase is about 310 stweek. Tne pups reach adldt
we:ght ar about 9 months.
ECOLOGY
>lore :s k n o w about h e e c o l o g o i ;oyotrs -Sar. ? e r h ~ sany other
carnivore IBeioiT 2 0 0 1 ~ ) Co"otes
.
o c x p y a vw,,ey of ilabiuts. inoicding g3ss;ancs. iescns, ma .mountam. ?ills? lo not icmpi:e Well
~ , r ihq c : cjriuvorcs xi x a y be hilcd by them or i v o ~ dareas a d
hablraz > c c ~ ~ . ehy
d these s?ec:rs. Studies $ a x documenred direct
m d 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 ' G m 1985;
Koehler andHomocker 1991: ,Murphy 1998)..4qoandP!erscher (1999)
documented behamoral chanses in coyotes after several years o i mcremed wolf a b u n h c e innorthwestem Montana. Paquet (1991. 1992)
reported that coyotes did not spaualiy avoid waives, bur actually followed their hails and scavenged woif-killed ungulates.
Inten~eciiickilling a..
m e m 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
~ a c size
k decreased !?om 6covotes to4 ia33% aechei. However. Gese
et al. (1996% 1996b) documented simiar annual vanations m coyate
pack s i e s 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, m d y
cyclic lagomorph populations, causes even greater m u a i 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 i e m i n environmental conditions (Peterson 1995; A j o 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; ~ & ~ e a and
n t M e n 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 t e 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 w i t h resident packs. they display a dominance
hierarchy similar to that of wolves (Camenrind 1978; Bowen 1978;
Bekoff and W e b 1986; Gese et a]. 1996a).Tne social orgamization and
land-tenure system mediate the r e ~ l a t ~ oaicoyote
n
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
N e h and Kath 19i6
B-n
1978
Todd et al. 1981
Pvrah 1984
Locsnon
Kansas
Texa3
.kesora
Ubem
AIbRta
Alberta
Babb andKmedy 1989
Montana
.exas
Colorado
Tennessee
O'Donagbue et ai. 1997
Henke and Brvant 1999
Yukon
Texas
~.
.Gesc et al. !989a
-
Denriv
(hdi~duahh')
0.8*
0.9': :.5-2.3c
0.24.AL
0.146'
0.46'; O X d
0.081)qdc
0.15'; 0.39'
0.99
0.26-0.338
0.358
0.01-0.09F
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; ~ n i w l t o nand 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; ~ i d 1985;
d
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 d o m a n t 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
as food re&urces become more
pack membersdisperse dming
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 s m c w e . h~ghersumival rates, lower reproduct~verates, h e r pack sue,
and!owerrecruimentlnw the adultpopulation i.4ndelt 1985; W m d m
et al. 1985; Gese eta1 1989%1996% 1996b: Widberg 1995; Knowiton
et d. 1999). Under h i g levels of exploiration coyote populations
!3skoi?;cc '&!Is !986; Grse , a d Grathe 13'11: C a r 5: d.;?o6ai T ,
concast S'rivlk e: ~ i(i999:)
.
reponed i change :n coyore ic:~nty ieveis
d w a g . w a r &at i.z not .le?e-dent on a zhange :n prey base. They
.heonzed jiar jic reJuced ac5nr:i !eveis heipea reduce : n e w expendue rc reduced coyor; ;ensir,, !ik;iy n respomc ro reducrd compt.5dimes ;r XI area ha linured food base. Coyotes
m p a c k 1esr
more and mt-ei less eduriig h e r months tban do coyotes iivlng as
tion for food (,bde!r 1987. 1096) ,,r brerdir.3 song younger females
(jLmw-lton er d. 1999).
m a t e d p x s or a c n e i B e k o 3 a d Wells 1981, 1386. but see Gese cr al.
lP96a). Large: :ne:-i
savings have been shown far i prepant fernale
p i e E~xpretmcy.Cayares J: c3pr:vity may iivc as long u I8 :,ears
living m 3 ;lack &an for a iemale living aniy w t h her mate (Bekoff
p i o u ~ gi951!, bur 2 wtia ?opuianons. 5fe sx?ec*imcy is considerably
and Wells i981). Howeve:, it is unknown mhefier Femdes livmg lo
shorter. >I&vaum 1Zes re?oned fcr w i d soyctss a e l j . 5 (Xsiiis in6
p a c k irc e a e r ahrepoduchvely because ili !he mrrp ?.raveddunng
%:th 1976), 14.5 (IGowitcn l9i3), and 15.5 yeas (Gese 19%)).
Drewaccy.
. - .
Cayores, sinuiar to wolves, are ;vpicaily rerrionai, md ?rimmiiy
Dens. Cayotes den in a vxetl, of placcs. iccluding brush-covered
rernaio mrhin thrr :e.nito~. However. they also a a y makc ecxmrzmsiopes, s:erp jar&. under rocx ledges. thicks:~, m d hollow lcgs. Dcns
ioriai movezcnts into nei&bnormg teAor.es. Dispersal from the nard
oiother m , a l s . ulciudins badgers :Toxiden i m s I, airc fiequentlyused.
site mliy be into a iacant or occupied temrop In m adjacent area or
E e m mzy have more !ha one enmnce (Bmison and Gliben 1985)
they may disperse lone distanczs (Xarrison er d . 19911. In general, it
a d ohen there are many hrercamec:in~runnels. Dcns may be osenred
is the pups ar subordinarc yearlings that typipicaily disperse (Knowiton
:o 'he iourh :o mar.jrize solar radiation iGier 1968; Hallen 1977; Har1972: Ne!lis and Keirh 1976; Berg and L3esness 1978: Gese et ai.
nsan and Ci?om 1982). The s m e den :nay be used iiom y r x to year
1989a. 1996a). Dispcrsal appears to be vaiuntaq a social m u nu*.D m i h a r a f occurs oniy rs;.ely (?jeiIis and Kdith 1976; Cmznzlnd
tional pressures inrensfi duing rhe winter m d avauability and access
1978). >Iavement of pups &Tong dezs i s v e y c u m o n (Hamson and
to food becomes more lhired (Gese et 31. 19963; Bekoff2001a). There
Gijben 198:). R a s o n s ior these moves n c nor known. but disrurbance
x e n o consistent sexdiiiereuczs m dispersal dismcz(Gese e:d. 19893;
(Harrison 3nd Giiben 1985) ind possibly iniesration by parasires may
Han;son sr al. 199 I), alr!rough in one study, female pups moved farther
he hcrors. Most moves are rtia"vely shar;; however, moves o i >? hi
than m d e pups (Xellis and Keith 1976). D~spersaiby juvcmles usually
are common iHarnson m d Gilben 1985). \ i n Wormer I 1964) reponed
thaf a maie coyote mcved four pups, ir.dividually, a distance of 8.3 :a. occurs during aurumn and early winter. rhougb same individuals do
nor disperse durx~gthe ijrsi yesr ma remain to prov~dec a e for f i t w e
T?x den and pup-rear,rz icn'rrties are the focal joint for coyore Pamilies
siblings (Kat:er 1995; Gcse et a]. 19R6a). D~spersalhc~renlonappears
far several months untd puns are iaizr and ;r.obzle ,.hdelt e: al. 1979;
ro be random, and pups w ~ lmove
l
%&I60 km (Bowen 1982; Gese er a!.
5zr;:son a d Gilben 1985; Bekodand Wells 1986; Haticr 1995).
l98Ya), with record dispersai o r 5 4 km (Carbyn m d Paquer 1986).
mnvity and Movements. Coyote:; xz acrive rhroushout the ;iv.but
Berg m d CC:iesness(1978j reported mean dispersil disrmces of 48 h,
rend io be mure :ic:ivc dunng the sariy mornlng and iround sunset
w&ich occurred ar a mean rate of d o u r I1 !m'week. Furays or ex(WoodmE mu Keiler l?;i2: .r\ndrit 1985; Ccse er 31. 1989b). Gipson
plor~roryrnuvmenrs xiso may occur beibre dispersal (Harrison st ai.
imxi Sealmdcr (1 9-: snowed a 9racrpal acrlvity p c ~ kat sunset and a
109 I). Increased morailty may be associated wirh & s p e d 3s h a i s
m o r peak at kaybicsk in .Arkansas. Activity patterns may c h m y sesmove into imimiliai xes and low-jecunty hab~tats(Tnikowsh 1980;
sonally (Hobman ei ai. l9923j or in response to human disrurbac: and
Pyrah 1981; Bekoffacd Weils 1986; Gese et XI. 198Ya). Observations
persecution (Grse et al. i984b; Kitcirn et 11. 2000a). Sczonal ac:iv??,
of coyote dispersal are summarked UI Table 21.3.
Fatterns d s o are obvious, sspeciaily dunng wmer months, when there
Home Rnnpe and T e r r i t o ~ .Home ranee
1s a c h a n g e b food b a e (BskoffandWells 1936: Gese et al. 1996a). For
- size o i coyotes has been
example, coyotes living in nonhen climates rest morz dmLng winter
smuied throu&our their nnge. Home range size varies geagapbically
ar.d sesonally (Gipson and Scaiander 1972; Be.bK m d Wells 1980;
months, when they are drpendcnt pnmanly on ungulate c a n o n fur
food t h a n d u n g other se~sdns,when they fe'eedmsinly on smali rodents
Laundre and Kellrr 198-l) and within a population ( S p ~ g e : 1977;
Bdwen 1982: Gese rt d . 1988a) (Table 22.4). vkiation in home range
( B c k o h ~ n dWells I98 1. Gese et ai. 1996ai. Hunrmg artemprs also desize D o n g resident coyotes wlrhin a popuiation depenh on energetic
crease durn5 winrer monks, when rodents are relatively inaccessible
b r x u s e of snow-covered p u n 4 one of &r mijor rodent food i t e m
requirements. season, sex, physiogaphic makeup, habitat, and food
(Linta yround si;uir;eis, Spemoghillrs annotus) are hibemarme, and
disebuuon (!!muore and Kdler 1984; Gese er d.1988ai. Home range
unplate w h e ~ ~ b ~ ltoi r?redation
y
incrr~seswirh snow acc.mdanon
rize aiso is inj3uenccd by social organizafion. Tiuslent individuals
,.ailv have 1 :iol;r.ge: 3 % ~scdc?r:. lower sum:yai, ,hc:raseJ num3,:; ,~i:iczling3rs?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; r z e 2.y mc:ese
-
T ~ L 21.2
E Average &stances of coyote disperrai movements from established home ranges
Juverulcs Orm)
Addts I h )
Sourcc
Wes
Femvles
Rooinson and Cummlngs 1951
Young 1951
Young !951
Robinson m d G m d 1958
bwthome 1971
Glpson and Sdander 1972
Nehs ma i(slrh 1976
.hdrews md Baggess 1978
Bowen 1982
Grse 2: 21.
1989a
h s o n :99?
'?vide p g s .
'Fernaic pups.
'Exciudes &ra 51one !c.mle hat moved a record dismce of 323.2 ion
Males
Fcmaleb
Both Scxcs
T-LE 22 A Mean coyore home range sizes from some representanve m d e s
Juverules and Pups ihm:)
Aouiti ilrm'j
Mdes
Females
Glpson and Scaiander 1972
Chesness and Bremickc: 1974
Maics
Females
Stare/Prowct
.*sas
.MmOmora
Mlnnesora
Utah
Okiahoma
Aibem
ManTexas
Gese zr d.19888
Windberg and Knowitan 1988
Texas
,Atbnsan and Shackleton 1991
Bribsh Columbia
Vernon:
Person and Hxth 1991
"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 r n a r h g and h o w h g (Camenzhd 1978; Bekoff and Wells 1980,
1986; m i l s andBekoff 1981; Gese and Ruff 1997, 1998; Gese 2001).
T ~ i c a l l yonly
,
pack members maintain and defend temrories; solimy
indinduals do not (Bekoff and W e h 1980; Gese ;ma Ruff 1997,1998;
Gese 2001). Fidelity to the home range area is very'high m o n g resident
an~mais,may persist for many y e a s (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; G a e 1998).
Coyotes are opportunistic3generaiist predators and ezt a variety of food
i t e m in relabon to chanses in availabiiity (Van Vwen and Thompson 1982; Todd and Ksirh 1983; b d e i t e: al. 1987; Windberg and
Mitchell 1990).Tney will consume food items rangng i
nsize u o n 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 ( 1 9 7 1 :oud rei.ior;ll m d sezonai diEerenczs in fee5nin.
habm oicoyores jiving in . k h s a s . ' h e most ; o m o n iood item from
168 coyore sromazhs were p o u i (34561,
~
p e r s m o n s (23"%),inseas
(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
F!cLw 11.3.Coyore feeding on elk calf carcass, YeUowstone Natianal Park
R-yormng. SOLICE:Phom by E. Gese.
:9--: 3 k o d I ~ . \Ve!:s
J
:Y8G), ' k t 3c:ui ?reiz~!onan ! q e ,xp:i:itss.
50th j a t x e ir.d tomesnc, ices o c z x :Gie: lGh3: I l i s i ! :%S-: i c s s
s.6 Crorl?e ' 905). In '*-xiomnn, the 3erccXue- :i;svate src;-,~cbsccn-
1978). C&on avruiabiliy dw.::g ~vmrcrpia~r
a h g e roie n coyote {oraeng m d populai!on e c o i c p :n n o r h c n rzxluns (Weaver :i):?: Todd
1'85; Gcse s:al. 19oh;i. 199hb).X;~roduiuc:ivs s m t u s n ~ y ? vd~us e~ m e
. .
:;.ore
fez5ng habirs. C ~ y c r :hat
~ s i r e provis;cning>r;;ls may ivxrcn :o
I x ~ e rmore
.
energecally '?ruduhiC' ?re? irems i s :omcared :o son:eproduc:~ve coyotes (Ti'd
'(;owlton !983; Hsrzscn u d Barison
13%; Sromiey 2000). Coycres in suburban 2 - s a xe s ~ x d l yadept
~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 u i o o d terns
n:clation to z c m i prey consmFnon cm confound:cs;ii?i mi should
b* considsrc? (tvexver m d IiorEma i979: Kelly uld Ganon 19971.
-
BEHAVIOR
Direcr abseriation ofcovote behavior :n d x ,wid is Jiffiait because
o i the slusive nature ofyhr species (Zsunan m u Bmdy 1975; Bskod
ZOOla). Howeve:. observarlocs of ;ovores in national varks iave orovided insight into many aspecs of :he bei:aviord e c o l o a oithe species
( C a r n c m d ;978; Bekoii' and Wdls 1980. 1081, 1986; Wells and
Eekoff :981; Gcss et al. !9Y6a, 19960, iYY6c; Gcse s1d Rufi- 1997.
1998: I l c n et al. 1999). Dam on behav~oraideveiopmenr x e 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 i r e only 19-24 Jays old (Knigh; :974; Bekodcr ai. 198;; BekoE
m d D u g a h 7 0 0 0 ) . T h e e a d y d e v e i o p m e n r ~ i i ~ r c i a ~ o nwithin
s~ps
litters a .o.p c m co iatirp 104.2 months liC~iehr
- IYlSi, bur the dominance
iurrvchy later m life may nor reflect the hienrchy o b s r m d w i ~ thr.
n
i n r r (Beko81977b; Knighr 1978). General behaviod p a n e x (postures, gcsrures. a i l novemmrs. facia! ~xprrssions,vocalkst~uns)uire
described m d 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, e x 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. m d Eisenberg (1973)
-ind K i c k a n and B n a y (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 u i 3e:secur:on
packs. is the donxl;mt idult h c t e r o s e . ~ lpair. 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 h e Sreeding pau and br-ome an d ~ u tah e selves (Gcse cd.i?%aa). These msocates (or 3etxanunsisl paruclpare
in : c t o n a l maintex~nc;m d puo i r m y (Harier 1995), bur out to rhr
sxrenr of the alpha pau (Gese m d R J E 11)97.;99F: C.cse 3001). Other
coyotes exist on :he landscape durside of *r rts~dentp n c b ar.d are
cnnsiderzd m i s n r or nomadic hci;.?dwds (Cme3zmd 1973; Bowe::
1982: Gcse c d. 19SSa). Tlesi: in-~lentsusmily m e ! ~ i o n eover 3
large area. i o ::or breed m d d>i 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,) m d :r ippe%-i :hat Ji3tr-sczs ietwezn ;:qc;tcs 1-d ,.~ciws=re
? ~ a t i t n n v rnthe: ;ha quiirar~ve.
,
.
!us; B .*v?h LI 'other S ~ ~ C L SS ~ ? ? c : e:;ti~c!~,
:hex ,> com;de:~ " l e.iar,abiiiry .kobserved sccrai oi:?vm~non (Li~wea1YX; Bckod
i r L .'!:?:Is
k:r;c-::!v
1980. is$:, 1C4h). '
a m w ires.soiirar:, :;diwduals sr:
cbic7e2,3::g md C'aesxss :?7S1 2utsiie 3 i h e iesdine
1?&31. :n 3th;: a r i q jr:i 1s J a c h c n ?.>is, W y o m : !,Canewad
. "Y
Y:E3esaffand'A~ci:s :?SO. 198:). md!asper,,%bem(Bowez:9;8);
zrcups ofcoyores a
: : ~ m o n i yobse~ied.?-ey slze maybe m p o m . r
in i5c:lr.g :nyo:e i o c i d i ~-/Isowen!YS:S e k o R m d We& 1980). in
,-i:puiariocs -.\-here the T q c r ire:, rtcms rhrou&our -be year are s-1
rodenrr, coycrssrs~dio he inpain o r x o s (3rkarimd We" i9Y6: Grse
e: 11. 1?8?aa).In ?n"ularions vhere lxce a m d s are mailzbie (a.2.. e k ,
..
Leer; s:tns: s live mdiv:duals or as w r o o . :oyores f o m 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 1pair or s goup j F ? h
m d Schwc~ner1979; Bour/er 1957: Gcse m d Grothe 1995). bsread
iooperxlve g o u p defense xppcxs to be the mqor seiecSve force favor:ng mc'easei soc:aii?/ ( B r z e r 1 9 2 : Bowen 1978; Lamprec51 1978;
B;kotTar@ SVr:;s 1980). However, :yen w i t h a resident pack, dominancs plays a !ai:e role in access to fezding on czrcxses (Gese et 31.
l"601 and niiuencrs thc abiiiy o i 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 r e s o w 2 iBowen 1978;
Cxr..e3zind 1979; Beioff and We!ls :980). In areas without larse ;in.
.
gxlzr: csrr!un, ! n c e ~ s e dpack slzz aiso occurs in :he brczhiing season,
which then f~iiiiurescaphlrc o i l x g e prey (Gese e: dl. i988b).
Predatory Behavior. There have been 2 k w 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, s n k pounce, head t h v c l o s e 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 ! q c r 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.: m u capture smallmammals (Weils
mdBzkod1981; hlilrrsy mdBoutm 1991; Gese i.1 I.1996~).Coyotes
generally hunt m I I mammals xione, even t h o u b t h c yack size may be
quire ixge (Gese 2: 111. 19960, 1996~).Cooperative hunting for s m l l
mnmm~iswould bc mrScient beczuse only one individual c m con; m e ;he prey ;tern (Bowen 1981; Andelt 1985; Gese :t d.1988b). It
h ~ been
s
shown expcnmenrIIy that coyotes iirpcnd on various senses
to ;ocarc prey. I11 order of decreasing hponance, they are vision, audition, and oifactlon ~ X e l l sand Lebnrr !gig), though these "pcorines.'
nay change depending on envlronmenal condinons (Wells 1978'1.
Pecbtirory behavic: mdhabiw ofwiidcoyotespreging on domssnc
livestock have received increased ~rtentionje.g., H e m e 1977; Shivlk
c t ~ 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
d i t e ~ c dcoyotes only 31.6?/0of the h e . Miem latency to attack was
-is?! long (47 mni, with consideraoiy vanabiiiy (swndard dewauon =
1 8 slir.) arnun_r amc.b. Mean kiilUlq rime, likewise, was cithcr long
(13 :nF), and cons~dcrrlblyvanable. Inmosr ;nstances, coyotes attacked
,.., ny biting the thros~and the r h ~ qdied 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 p r o g m . In xenerd, chan~es
in dnixal ilze and behavior, differences h s o u p cohesiveness, soclaiity, grazmy kspersion, attmnveness. and m t c m i protection ail adefecI
v~inerabiityto coyare predation (sez re~nzwby Knowiton st ai. i99Y).
R->.dbrrx 2: A.(1997b) and Bromiey (1000a)d o ~ m e n t s dthat coyotes
~ r x n i i y'oiled rhe m d l e s r d r !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;
E r a s : 370:Zcwyer !987\. lV?ea ;oycies mi.!;large -aid ungulates.
e.lvusrmsnE! ?ac:~rs sue= 3s ;now de-jrb and 3ardnrss o i the zrust
..
2 2 i r i ~ o r m ::l! the I I I C ~ ~ Siir
S :xure :f're mex?t iGrs; m a Grothe
,995j . ?:eseacr ar' h e i p h a ?ax also appears to be i m p o m 1 i
n 'be
outcome oithe attack. Generally, younger anlmds do not pmicipzrs in
the atiack sequent:. It appem that the m b e : o i cayores m the pack
is not as imponant as which coyotes are involved m the araci. Even 2
a pack ofsevec coyotes. a d y the alpha pair was hvoived in attacks on
1aree;inpuiates IGese andGrothe 1995'1.Also. the abuiwoi~uneularesto
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
t g e s 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) d s o influence coyote vocalization rates. Howling amang coyotes
plays a role in terntonal maintenance and pack spacing. It s i - 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
es
the majority
and Ruff 1997; Allen st al. 1999). ~ l ~ h a c o y o t perform
of scent-maridng duties. Rates of s c a t 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 conblbute 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~iedterntones also mav be
communicated via scent marks.
MORTALITY
~MortalityRates. 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 ( c l 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 K e ~ t b1976).
in \iellowstone Nanonal Pxk. prevalence of antibodies against cam e j m o v i n s (CP? was 10046 for sdults, yearlings. a n 6 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
o i the coyotes bad antibodes against C?V m o m a s et al. 1984 [n =
11841; Gese :t d.1991 [n = 721). In Georgia, 65% of i7 coyotes had
antibodies against CPV ( H o i m 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'~0diesagainst 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 v i m ; Guo e: al. (1986) reported 56% of 228.
W~lliamset ai. (1988) reported that 5096 of 10 coyotes in Wyomng
tesred posirive for this w u s . In G e o r ~ ano
, coyotes were found to have
been exposed (Holrman eta!. 1992b).
h e ~ l s n c oe f c a k e inkcnous hepatitis virus antibodies was 97%,
82%, 54%. and 3354 for adults, yearlings; old pups, and young pups.
iespecnvely. inYeUowstoneXationai Park (Grse d a i . 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.
et d.
. (Gese
,
* old nuns.
. . , and .voune .ouns.
. . respectivelv
.
1997) T h s !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
mav become infected with Y Destis bv
and Huehes 19921.
, Covotes
,
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 ~ l a e u e(Bames 1982). Changes
- in prevalence of
plague h the coyote population likely reflect prevalence of piague in
the smal-mammal Drev base.
-
. .
&ely low levels ( 4 5 % ) ( ~ e s ezt al. 1997). In Texas, Trainer and
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, H o h a n 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) A d 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 lepCauses of Mortality. Most shidies indicate that human acttvity is in.. . ~ .. ~ ~. : : n ~ . l ~ g ~ . ~ r ~ ~ c ~ r ~:::vo!es
c ~ ~ ~>dv!sd~
~ ~ ~ m e1981,
. l c ~ ~tospirosis
~ , ~ i ~on
~ coyotes
u l ~ is udnowo, but infected dmay survive and
caniers
for a short time.
remain
I : . ; : 8 ,'
r 2: I 1
,s
! i . :3h9a, Mmdbun:
Coyotes aiso cany a M n e ? of parasites. Ectoparasites com1993, although preiationhy large cmivores (Peterson 1995; Aqo
monly found amang coyotes include fleas (the most common external
1998: Crabtree and Sheldon !999) and stamanon dunne crashes of
parasite), various ricks, mites, and lice: see Gier and Ameei (1959)
food resources also may be substantial m o d t y factors. b~seasealso
and Girr et d.(1978) ibr species identiiication. Intemai parasites incan be a factor. especially among young or the year; pmovirus ialled
clude several s~ecieso i flukes (Wematodes). tapeworms (cestodes),
several radio-marked pups in ~~eilowsrone
iGese et al. 19971. Even in
intestinal worms (nematndes. ascarids), hookworms (ancylostomlds),
5&tly explo~tedpopulations. mosr mortality is armburable to human
heartworms lfi!aroidsi. eso~haeealworms (spinxoidsi, lungwarms
causes. Human exploiration can be substanr.d in some coyote popuianom (&owiton :9?;; &owlton et ai. 19991.
Diseases and Parasites. Ssrologicd analyses for anrioodies in coyotes have shown - b t they have been eerposed to =any diseases.
.. .
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;,,, n a y m p ~ s b ! e sm a mA:d ~siliferi o m crrrdiovascuix- &sexes. 30:Ju mesrjsiis iT>cmtcn ;r a.I?YJ:. mange (Pe3c-n27 21. :!?El: ?PX=
'yVx~ibe:g I?OJ'.. m u c a n c x F.:r ie\'?ews m a :cxcncmic mdyses
dis x t ~ acc
~ lmemii. 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~ 1 3 ~ 2wlrh
6 tcorh weir (Gier !968). Before tooth sec:ionmg for cc; n c n t l i . anaiysis; 3qc class ijui-exlzs m u adults) can bc &srrn-gushed
basrd on the r:!ar,ve zuip C 3 V l v size usinq miiopaphy !,LKnowitonand
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. m u !enmh
. ofthr hind fool iGier 1968: Bckoff and lameson
19-5: B m m ;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~ousbertieen wclzht and age for !&3O days m d 3 ;-15.i days
x c 0.999 m d 0.995. respectively (Bamum et 11. 19-99).
-
+
-
+
ECONOlVEC ST.~LTUSAYi) hI.LU.\GEMX.W
Coyotes arz ,ncrlms of success. By t a k n ~
ldvanrage of poorly devised
dorncst~car,unorscrlces 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 c u i h r ~ tates
l
as well as to piov:de food and fiber (Knowiton st il.
1999) DiiTerenc.5 in group cohes~vcncss.sociality, g a x n g 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 ~ l coyores
l
:GI! sheep. Some are never exposed
to shecp (Wagner 198Y), whereas others do not develop sheep-hilin~
rendenc:es jLr.S. Fish m d Wildlife S e m ~ c r1978; 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) m d Windbcrg r: d. (1997b) Eoorrsd a hi=! numnbcr oicovorss thm killcd and consumed !ivcsrock. In
the internountab West. Till and b u w . t o n i 1985) m d Bromiey (20001
dzmonsuatrd ttut aduit coyales wirh pups were more lkeiy to h i l iambs
thanadults \i;thourpups. In dunnzsr. renironal ioyotes innoA-coaswl
CdiLiibmia laii I m b s as soon= h r v nbccamc~vailabie:nDecember and
January, oursldz the p u p - r C d ~ gs r s o n (Connrr 1995; Sic!ti 1996).
Economics of Coyote Predation. Tne m o u n t of :ivestock !osses that
produce3 imibure :o coyotes is J sonrenrious issue. Wildlife protrcnonsdvocztes : i a i m ~ hie.u
t or nu depredarions actuaily acc.~,where=
;rroduce~skdicare !asses From cayates threaren their economc livclihood !Caine m dl. 1973: Ge- et 31. 197'; CS. Fish md Wildlife S z M c e
i?7S: Wagner 1988; Knowiton er ai. i094). In iiruanons where here
was prehror :onnoi. losses of sneep ro cayotes were : . i L ' . O ~ o far
I m b s =,ld 0.1&2.0?/0for ewes (US. Fish and Wiidlife Semic: 19781. h
%>ward 19-6; 3rawiz:i I?--: Heunr 1 9 7 :vllmoz 1977: Llc.idoo and
:ie!:e~o\v 19-8; O ' i 7 ~ re: ii. :!JP>
In.:?%. :he zcanomic value
ui domes::^ sheep 10s; ro pre&ron was reporrsd is $17.7 .milion
jf >.rri.-!rure 19q5i. ie=irr m 6 S:;m&e (:978)
m d L~.S
De3m:s::: oiS=$;?liture i:?95! r n a n z d - h r c3yotes w c ~ e
rsspc n>,o:c $1 h e ;lajcr;.i :i 111 she:? lost :o p r e r o n . iiowe.ie:, ;r
,
is hpocjr.: :o corsiccr :hat coyctz prehnon is ilot 'ie najor c 3 u e
o i :css;s Cis. Fish ma 'Xildiiie Serilce !9-8: Bekaii 1979b). Eariy
s7 1:. ( 197.1) found thzr -7% a i the rotai cc3nonic ;lass o i iherp k
:3-%1?7l was Lue :a Asease (43?'0), w.spec:fisi zauses 131?6j. m d
pre&rcrr C3"/oi.
11-5. Fey-z-r
Depredation \.lsnszement Protecting 1ives:cck or wlidife species
i o m :oyores :ccn;;s
;ons~de:anon ut';qal. soc:d, sconomic. biolosicd. :;c:m~wi.
5:bcal fac:ors (Stener an3 S h u m k e i97S; Wade
13-5; Bekoff 1908. 20Olb; fficwlton et ai. I'J991. Succrssliii resoiut:on o f c o d i c : ~should cansider rhe efficac:~. seiecuvlty, and ejicirncy
ot' vanous proc:dures. S;.re.al t e - . k i ~ e s x e avaiiabie ?or reducmg
czyore depredano~son !ives:ac!c (Siemer m d 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 c o ~ n o in
i either a preventative or ccrrecnve context. Procedues ;Oat are more b s m p 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. S z v e d rec!mio,ucs have been used to detsr
coyores %om attachnq livesrock (Linhan 1984a: Wagner 1988; Fall
1990; Green et i l . 199.'). W b e r e x 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 g a m e s , s y n c k o m ~ g
binhmg, and kcepmg young m a i s 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 o f u are.>
~ !Ori d e s t a 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 s h e r ~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 x e used 35 !ivesrock guards jLinhm. e-Ld.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 m d !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 1 3 7 ; Bums and Lhson 1997). investigations of
condinonedmste ,lverslan usmg iihiwn ~ u o c d showednuxedresults.
e
Some xscarcherr reported success arrcducmg food consumpnon (e.g.,
Gustavson st xi. 1974. 1981; Ellins m d 4 1 m 198:; Fonhmm-Quick
e: 31. !985a, 1985b), whereas orhers repaired no ieduction oi'predation
bv. couores
iBekoE:97i; C-nover 5 ai. 1977. B u m 1980; !983; Bums
.
and Connolly 1980, 1985; Bourne m a Dorrmcz 1982).
Sttemats to reduc; covote reproducnve rdres and popuiabon leveis ]lave been :nvesr;gateu with rhe a s s u m ~ c o nthat iewer Gayotes will
rzsuit in fewer i;e3re&Iions {Knowiton sr d. 1999). Copre repraducnon ,was reducea'sing diethyistiibesreroi [Baiser 106i; L a b a n cr ai.
1908:. bur was :~mcmcncdwithour effcc:ive bair ieiive? svstems.
.Qternar.veiy, reoroaucrlve inrede:ence usmg jret51iz3tion 3f temtonxi. breedin?:
bem , - ~ e d l v ; ~ ~ a v reduce
:o
de~rehtionsby
- ca\otesrn;lv
.
coyarcs ~ B r o d c y2000) by i b & g ;he pre.&rov bciavtor of adults
when nro\.is;orim!g young (Tii1 m d ~ k o w i t o n1983). With respect ro
chernosrerianrs. Sotin rcqianon ifedcrai w d S ~ I iS2nd disnburion x c
prohemrn:c , SteiXug 2: 31. 1978).
Cz,yote Remuvd Techniques. X%7senncnrzmcwi :ccimli;ues do -or
stop deprehcioos, rmoving one or more soyores may acheve
mananernenr
o b i e c 5 e s m o w i t o n e: ai. 1999). e s ~ c c i a i i vii the
removed anlnais are the "problem individuals" (Linnd 5: a 1 1999).
Removlnn one o r w o :n&vudi;als m a amall area ii.e., corrective c c m o l >
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. s u e of the a r e a 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 c a h g and s h o o ~ can
g 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 r e m o m e coyotes
.
responsible for depredations is the use of the livesrock protection collar
f M c B n d e 1974, 1982; Burns et al. 1988, 1996; Connoilv and Burns
1990; Connolly 1993; Rollins 1995). These devices reiease a toxic
c h e m i c d 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, a n d M 4 4
devices. but tbe selectivity for Che offending coyote(s) is lower (Brand
et ai. 1 9 9 3 . Most depredanons are attributed to terntonal, dominant
coyotes (Till andKnowlton 1983; S a c k 1996). W i t h t h e s e tcmitorizs,
coyotes are less winetable to capture devices because of thelr farmliarity of the area ( H a m s 1983; Windberg and Knowlton 1990: Windberg
19963, which makes removal of offending lndividuds d f i c u l t (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 s a f e q of pilorr and gunners, seiect~vityof anunals, aod economtcs
p u t 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 o r predation
on iivestock needs to be re vented when more b e n i m tecbniaues have
been ineffective ( ~ n o w l t o net 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). As noted previously, before any popuiation reduction measures are impiementeii the technical, legai. social, biological, economic, and ethical considerations as weil as the e5ciency,
effectiveness, and selechvity of the rechnique shouid be considered.
LITERATLRE CITED
.h&wvs. R. D.. ma E. K. Bosgess. 1978. Ecaiogy a i coyotes m Iowa. rases
21965 in M. aekorf ed. Coyotes: Biology, behmar and managemcnr.
Academc Press. NCWYork.
.Qo, W. >l. 1998. ?'he effecr: of recoio-g
wolves oc coyote popuiaoons. movcmenrr, behaviors. and food habm Dissemnon. Umven~ryof
.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
h s s . irhaca, NY
A t h s . 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. Evoiunoo of the cerebellum m the gmus
Cans. Journal ofMammaiogy 52:9&107.
A h s o n , K. 7 ,and D M. Shackleton. 1991. Coyote. Conrr iornnr. ccoiogy in
a rural-urban enwironmmt. Canadian Fieid-Nadist 105:4>54.
Babb: J. G., and ,M. L. Kemcdy 1989. An estlmate of *urn
density for
capres in western Temessce. Journal of Wfldlife M q e m m t 53:18&
88.
Balser, D.S. 1964. Management ofpredatarpapuiatiom withaatifeidIity apenrs.
30-d
of Wildlife M y w e n t 28352-58.
Barnes. A. M. 1982. Swelllance and conmi of bubomc plague in the Umred
Swtes. Symposia ofthe Zoological Society (London) 50:Ui-70.
B a m q D. A.. J S. k e n , i T Flindcis, and N. L. Gates. 1979. Numoonal
levels and growth rater afhand-reared coyote pups. Journal ofManmalogy
60320-23.
Barrene. C., andF. Messier 1980. Scent-markingin hee-ran*g coyotes, Canis
lorans. Animal Behaviour 28:81&19.
Bekoff, M. 1972. An ethological study of the development of social interaction
in rhe genus Conis: .-I dyadic analysis.Dissenation, Washington Uluversity,
St. Lollis, MO.
Bekoff. M. 1974. Soc~alplay and play-aolicihg by infant canids. Amencan
Zoologist 14:32341.
Bekoff. M. 1975 Predation and avenive conditioning in coyotes. Science
187:!096.
Bekoff, M.1977%Conis inrans. Mammalian Species 7 9 - 9 .
Bekoff, M. 1977b. Mammalian dispersd and ontogeny of individual behavioral
phmowes. American Namdist 111:71S32.
Bekoff. M 1978.Behavlad development in coyotes and eastern coyotes. Pages
97-!26 in M. Bekoff, ed. Coyotes: Biology, behavior and management
Academic Rcss, New Yark.
BekoE, M. 1979a. Ground scratchz by male domesic dogs: A composite
rimal.
Jaumzl of Mammaioev 60:84718.
u
Bekoff. M. 1979b. Coyote damage assessment ithe west: Rtview of a repon.
atoseience 29:754.
Bekaff, M. 1998. Encyclopediaof animal rights a n d d w c l f a r e . Greenwood
Wesrport, CT.
Bekoff, M. 2001a. Cunningcoyores:T i l e s s tnchrers, protean predaton. Pages
381-107 in L. Dugat!&. ed. Model systems in behaviami ecology P+aceton Umversiq ks,
Pnncetoq NI.
Bekoff. M. 2001b. Humartcarmvore interactions: AdopMgpraactk srraregies
for complex problems. Pages 179-195 in 1.L. Gittieman, S. M.Funk, D. W.
Macdonald and R R Wayne, eds. C m v o r e c o n s e m o n . Cambridge
Urllvenity Press, London
Bekoff, M., and 1. Diamond I976 Piecopulatory and copulatory behavlor m
cnvotes. J a m 1 of Mammalow 57:372-75.
~ e k o f f , . ~and
. . L. A. Dugatkin 2000. Wimm and laser effecrs and the development of dormnaoce in young coyotes: An inreganan of daia and theory
Evolunanary Ecology Research 2871-83
Bekoff, M., and R. Jmieson. 1975. Physical development in coyotes (Cnnrs
i n m ) witha cornpanson to other canids. Journal o f M a m d o g y 56:685
-2
Adojan, A. S.; and G.B. Kolenosb. 1969. 4 manual far the idenniicauon of
h a k of selected Onmio mammais ( R e s d Report 90). Ontano Department o i h d s and Faicbu.
Allen I., IM.
Bekoff. and R Crabme. 1999 4n observanond siudy of coyote
(Canls lonow) scent-marbg and tmtonality in Yellowstone Nauonal
P a d Ethology 105:289-302.
Andeit, W.
1985. Behavtoml ccology of coyotes in south T e m . Wildlife
Monogrsphs 941.45.
Andeit. W. F. 1987. Cwote~r&nan. Paces 12840 in .M No& J. 4.Baker.
Bekoff. M.. and .U. C. Wells. 1980 Sacid ecology and behavior of coyotcs.
O n m o Trappers .issaciatian
Andelt W F. 1996. Cmvores. P y c s 133-55 in P R Krausman. r d &geland
ulidlifc. Society far Ranse Management. Denver CO.
Andelt W. F.. D. P. UthoE. and P. S. Ginson. 1979. 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:.
Sc:ennfic Ammcan 242:13C48.
Bekoff, M,%~M.
C. wells. 198l.~ehavlauralbu&edngbywjldcoyotes: The
ia&xncs of food n s a m e s and social a ~ m i z x t i a n h. i m a l Behaviour
29:79&801.
1986. Socral ecology and bchavlor of coyotes.
Bekoff, M., and M. C.
Advances m the Studr of Benavioi 16:251-338.
Bckoff. M.. H.L. W. and J. B. ~Minon.1975. Behavlorai manomy in m d s
by discruninant iuncaon analysis. Science !90:!22>25.
BekoE. M..bi.TyrreU M,V E Lipen. an6 R A. Jmieson 1981. Fighting patterns i
x young coyora: hitiaaan, escalanon, and assesmem. Agggcss~ve
B e h i o r i:Z2.W.
0"
I..T.>I. %I-e. l c d I 41. Z c g h m . 3 % 2-ccnjiiion
&~:ezces
;jYarc (Im!r:nrrnrui.ii;w!ing. i m c n c 3 n L t i d l a n l l S ~ ~ i i.30.111-is;
,.,3:
W 5.. ma h A. C.csress. : P i 4 i z o i u g .>i;a)orss n n o r 3 e z
iinr.inr.sam. ?=ees 1:0--17 :n M. Bcrof. $3. Cayxes: Bioioe. beenwar
m d nr-agemmi. .4wilcrmc E e s s . Yew Ybrk.
3crge:. J. :O-3. '?raaior imssmmr' LS I liefensive s m t s g . . a e n w n Mid!and Y d l s r :Oi:!9'-39
3 : n i r H. I..m a 1. 5. Green. 1081. Na>,a!a ise i;nu;c'-bred doqs for man:qcmem a i p r e u r a r j . Joumai oiilaog? Li-qemcnt
j8:i 1 1 5 .
~ b i a i z y i.
, 5.m a I. C. .LlcGrew 1984. Dt2s:ennni uinsracCiq ~ i 1 a m o r:a
cayore ?i;&non. ippiicd ha! Benancr Sc:mcc 1::34%5;.
aaggsss. E.K.. a d % X. Henderrm. ;?X. Rcgonni rve;i.hts u i b a s coyares.
Trriacaoos af!he h s a s Academy o i Science 8O:?LdO.
Bourne. i.mdLL J Dorimcr. !982. A
. field resr oilitlxun ;Sonde avcrslon ro
rccuc- ;wote ? r e u n o n J n domesiic me=. Journal oiWildllic I m a g e r n e s 16.235-39.
Bourn. 5.. ind B. D.Cluff. 1989. Coyote prey cholct: Optimal or o p p o m i s u c
for~gmq'?.A c o m m m r journal ofW-iidliie Management 53:dbZ-66.
Bowen. W 3. :?i8. i o c : i oqmizanan ofthe coyore in relmon :o prey size.
3isssnaoon. Umversiy a i B n a s h Colllir..oia, Vancouver, Cmaw.
5uwcn. W 0. 1981. Vaistian in iayors snc:al aqznizanun: The :duenc: o i
?re:, sue. Canadian : o w a i oiZ~o1og-j59-039- 52.
Yo\ven. W. D 1452. fiorne :mpe and sprnlnl orgm::woon oicoyates m Jasprr
Nuiiinunal Pui.41bcr;i. joumdi of.WtIdLte Managemnr 16:10i-15.
Boucn. W D,and I. M. C o w , . 1980. Sczzt nanung h coyores Calnadtan
Joumai o i Z o o l o g 58:173-SO.
R o y e r R. T.!9X7 Cyarc group size ieiative io predanon on trrule deer I s m maiia 5!:j15-26.
Yoyd D . m d B. W. O'Gara. 3985 Caugar?iedai:on oncilyatrs. LLumcIer 66: !7.
Biand D. J. N.?w~il,
and W kt. Scorf. 1995.
mriuencc oi:c;&;irrcmovni
ofblack-backedjachls on the .;ficicnuy of Goyore gcuers. South A k c m
J o m a i oiWi1dlife Research L S : W 8 .
3mw!ey, K. C. 19":. D a m e d c siirep r n o n i n y dufmg n d after :eso oisevrrui
predaioi control methods. Thesis. 1,:niverjly o i b l o n m a LLissaula.
Brom1e:j C. '000. Cuyotr stcnliurion 2s 1 mcms ,of rea.a;mq predsnon an
domesr!c !ambs. T x s i s . i;tm itlrc ;wveniry, L;gan.
Bnms. 5.H 1970. Winter prc;lat:on ,of yalaen c q i r s 2nd coyores onpron+horn
u;reic:pes. Canadian iicld-Narunlisr 34:;0!2.
Bums. R. ;. 1980. Evaiuaaan oicondinoncd ~rcbanonaversion for controlling
coyote predation. J o u d of Wildliic Llanagcment C 1 . 9 3 W 2 ~
Sums. R, J. 1983. .Llic:ocncapwiarcd l i t h ~ mahiundc hut avivers>ondid not stop
coyote predanon on sheep. Journal of Wiidiifc 4lminagement 17:101&17.
3urns. R. J., m d G T. Cornoily 1980. Lithum cNonds baa averiion d d nor
injuexce prcy lulling by coyotes. Procecdingr of the Vmebmte ?:st C.,nferenc:: 9-:0&204.
B u n s . R. J.. m d G. E. Comolly 1985. A iommenr on "Coyare conwal and tvtc
aversion." Apprnre 6.27&81.
Bums, I<. J.. m d J. R Mason. 1997. Effecnvcncss aiVlchas nan-ierhal callas
In dcr*.mnp ioyorr attacks an sheep. ?:oc;rdings o i ;hi. '+nebme Tesr
Canimmcr 1 7 : 2 0 t 6 .
S m s . R. I.,G.E. Conn<,ily, m d P. L Savare. 1988. L a r g iivesrack prarrc:Ion coilas criecrivr q m s t coyores. Oroc:zain;ls of the 'genebnre Pesr
Canrkrence 13:215-19.
Burns, X. J. D. E. Z:mlicka. m d P. I. Savxie. 1996. Eiiecuvmess of !we
livenaca 3rzrecnon ~ o U m
a s m s r droredanne- sovotcs.
Wiidlite Soc:cri
.
Builetm 34:!?3-27.
C ~ t w J a eY
, ii. 1947. .A dcrrioyare episode. Joumai oiMmmdogZY:3&39.
C ~ m r .S. .A, J. .4. Xxdlec. D. L. Allen, R .4. C~uley.M. C, iiomocker, A . 5.
Lzspoid m d F. H. Wagner 1972. ?redator conrrai--1971-Repon
ro the
C ~ u n c i an
l Enviranmcnmi Qiuiicy and the Dep-rnt
oithe inrenor by
rSe ;iliv~rocyC o m m i ~ e con Predaror Canuoi. Colmcii on Envkonmerrul
Quaiiqr m d L-.S. D c p a m c a r dthr hrmor. Washmgon. DC.
C:m;.limd
C 1. 1978. B e h a m o d c c o l o g oicoyotcs anthe NananalElkReNgr,
J x h o n . Wyoming. Paces 267-9.1 in Lf.B c k o d &. Cdyores: blolagy, Behav:ar ind management. hcademc Pxss. Yew %&.
C x b y I.
~ N.. a d P C. Paque:. 1986. Lanq dismc: movemrmaia coyare kom
.Zdmg Lioumin Nanonai ?azk Jom.d oi'Nildiie Managencnt 50:39.
Chsnens. R, A . m d T . ? Y r m c r c c r 1?7.l:<uacrmge. trrnrorali~~,n,dsosacu'jiiiry d i ioyol\/oresin nonil-scami Liimcsou. Cfiyore Rcsexci Workshop.
Demer, CO.
r;?. f 'V 19-?. Iniinence ~ i j 3 ~ : ~ 3 b de:uiy
bir
on :oyovors ?apui3oon ;hminge.
Jaumal ii7Xiidiie Liminag;.uezi 36:223-Ic.
;?nuer (2. L . m d i . hi. L;ve!css. 10-8. ilr.::er .:i:hec;ll.oie C z n w ,'amam)
m xzscx L:m. :ow.ai a i T i i c h i e Ztsz.:es i1::4749.
C;rzc:. \I. ii. :9?: !denr;fJing a r r e z t s .r;ayere ,reanan an iherp jn i
nunhem C a h o m a m c n . T~esls,Cmue:s>y of CalIrima. Bz:kc!c.i
3 e 2 a e ~3
-
-.
C.i~m;r 4!. M..M. M.!:lee: 7 J. W-cilcr. m a 2. 3.XlcC~ilou@.:go8 r:e-,c:
o i ?avore rc:noval >r. :nee; 2ezreilar:an a lorze~
<dirhma. lo&
ai
W i i d i e Manaermen: 5;:5?1&39.
Cjnni:;lr i.E, 19-S. Frrdrzi ;ooa;.i ma cayore popuiat~acs:.Aren:cw aisimulmon nodsis. ?~gesX - 4 5 in M. Seiorf. sd. CJ~OICS:
B x o ~ oY~, &, ~ ~ C ,
m d maagemen: .Ac>dealc ?ress, Ncx York
C~nnaily;i.E. 1945. i m d i hrnung taker jhces-&lling coyores n wsrm
Lionrma. LS. Farest S c r l c r . Ccmrd Tecmcai Kc3oit &\I ! 1 4 1 8 t
88.
Cdnnodv, G. E. : 9 9 ? L.vcircci prorecnon : o l l ~ sa &c TJoiieo Starcs. 19881993. Prcczcdings o i r$e Grear ? ! m s W i l a h ~D
i -e
C~lniroiWorkshop
,
.
- ?
.-
1 ,:22-:,.
Cannoily, G. E . and R. I. B m s . 1900 E5cacy of compound i080 iivestock
prorccnan cnllm? for hlilnq coyotcs rhai i m c k s a c q Procrrding~o i the
Vcnejmr; Pesr Conierence :4:36%7b.
C:maUy G. Z., R. LL. T h . W E. Howard ma W M Langhursr. 1976.
S h e q hiling bcnavioi o i a p n v e coyote.. Joumni oi7YiiiidiifeLPmagrnmt
W:1007
Canovcr. M. R, J G. Frsuck. mind D. E. \.lille:. 1 9 7 7 inexpenmena1 cwluanon
afuslnghstc ave.s:on :o cocuol sheep loss dueto ccyore pre&"on. Journai
ofWilfife Lianneemenr 11:775-79.
Caolahaa C. 1490. The usc of dogs m d calls to : k c cayores uaund dens m d
iesnng ue:w. Proceedings o i t h r Venebnrc ?err Comkrcncc 14:260-62.
Cappinger, X..J. Larcnz, a i d L. Cdppinge:. 1988'. huoducmg iivestuck g m ~ d m c
di-s io sncrp and joar productn. Procecsingr o i 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., m d i. W Sheldon. IP9R Cayotvotes m d c a d Zaeristcncc m
Yc!loiustonc. Pngcs 127-j3 in T.W Clark A. P Curie:. S. C. Llmti, m d
F . h r : m . 5"s. Cmivores n ecosystems. Ydc L~nlvcrstpPress, N e w
Haven. CT.
Crooks, K. R.. and M. E. S o d = 1999. Mesopredator reirasc mi avliaunal
txtmcnous m a hgmsnred aysrem. Nature 400:563-66.
C m m g h m , V 3..a n d R D.Dm'or*. 1970. Rcccnr coyorencord i r o m F l o n h
Qu;irre;iy Jbunai d f h c 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 k t foxes. Jo-d a i M m a i o g 79:2OL211.
Davisou. R. P 1980. The c l i k t of cxda~rationan some pardmcrerr o i coyote
papuinnoris i)~rsmarion.Utnh S h t e Umveniy, Logan.
M. G. Cropsey 1978. Fiela test o i a eoyoie~prooifence.
Dc C ; i l c s t ~D. 5..
Wadlife Sucieq Bullern 6:25&59.
DeLorenzo, D. G., md V W. Huward, ir 1976. ivaluanao of sheep Lasses on
a r a n g l m b m g openoan wirhour predarar canzai .nsourhrssrern New
Mex;co. Find rtponza :he L.S. Fish ma Wild~feScmicc. Denver Widlife
Rcscarch Cente; New Mexlco State Umveniq, Laa Crwes.
D ~ c e ,L. R. 1942. A i m l y of dus<uyore hybnds. Journal of M a m a l o g
23:186-92.
Drcwek. 1,ir, T. H. Uoun; K. J. 'finurnan. and E. i. Bic.helL 1981 Scrolodc
svldencr ai1eprospaosis m a r o u t h ~ m ; \ n z o ncoyore
~
popuiat~on.J o m d
ai'hTidlife Disessrs 17:33-37.
Cads, h 3. !948 Ecropmsltes iiam a scnrts ufTexas coyores. J o d o f M m m a l a g 29:26&7i
Cxly, I. 0.. J. C. Raethcii, md 0. R. Brewer 1974. .hccanarmc study o f
p r e c n o n h the !dho m.gc sheep m d u s ~ ,1970-71 pioducjun q c l e
iPraeress Rsoon No. 182:. Idmo h u n r ; i t u d Rescmcn. U m ~ e n l r io i
idah;, Moscow
E!&r, 'N R., mnC.LL. i i q d m . 1977. Us;of&sc~mmrhuncuonmraxonomc
ie:;.manon
o i cmds t o m Missow.. Journal ui Mummaiac 2!:17-
-
ilii. 41. W ! 990. Canno1 o i ~ o y a t cdepledation an livestock ?regress in resemh m d dcr,eiopmenr. Proceedings of rhc Venebmre nest Cooference
143-51.
M. 1999. T I C oicmare survivor Audooan IOl:5Z-S9.
Fisher. J. i 9 i S The pialns joy moves cast Nanooni 'Niidbe 13 ! L i 6 .
Fonhman-Quck 3.L.,C. R. Gusravsan. md K. W Xusitmk 1985a. Coyote
isnuol and msie &yesion. p p e n r c 6 : 2 5 3 4 a .
Fonbmm-Quck D. L.. C. R Guswvsan. and Y W. Rusmak. 1985. Cayores
m d 135:~ 3ven10n: 7 3 m~bors'
~
. q l y A p p r ~ ?e : l S L 3 0 .
FOX.>L. V ! Q 7 O A iar.~ar~rive~:u(Iya i a ; ~sveioprr.mmai5c:al ;xpressions
m zamdr: 'Yoii. ;ayore. foxes. Br5~iv1or;6.19--3.
i-ale:. .: K . m d L. 3. Kt:&. 198:. Non-.>\en~npnsmues 2 i cayores m d
solves in norrhmstern .Ube.m. J o m d a i M a m m a i o g 62:403-5.
S;&n!,
:??6 5i;:nenrss
ai-iainni; r o e z m a l m m n r r ar redusag egg
~;n?unipnoa iy x m a l i m pemron: in :xpermencd maiysis. *Icsa,
~- t r Si;is
n
Univernv, h g m .
Zanl. j 'T.. ma P V L:hs:.
: ^ 5 . S i ~ i o p l ci r n a n i ~ r in
j the coyare ( G n u
Icinrm:. loEsi d f i C $ m p m ~ v ePhysirriogy m d Psycnotag 39::070-:6.
%ouston. 3.3.19-3. Fk IS .Inre:-spch:i ioad k r - m o r e s m norrhern k
!:owsroiie Ndnona Pinr. ; o m d oi-lppiiei E c o i o g 1 5 6 5 3 4 1 .
Gowars \V.E. 1944 4 mrms :o iisnngusn j r d i s a i zcyores m d 2omes.c
mu,- J c m z l o i L 1 u n m ; U o ~50::03-7:.
lacdon. 3. H. T 195:. ? n 3 . C:assuicztir,on ofrhe s c r s dicayores. P q r s 237341 : i z 5. Young mu 3.H. T.Jackson. cds. X,r cieve: sayare. 'k~&fe
l a n q c n c a c !mnsiimze, INashmgron. DC.
:(;!y.
3. T , md E. 0. i m n . :99:.
2fec:s of prey size, meal rlzc, mesi
cornposioan md dady fiequenq a i < d i n g dn the recovrzy o i mdenr
rernsm iron camvare sia;s. Cainndzan I o u n a i o i Z a o i o g 75:lYIl-iKcnneiiv.
,. J. !. :97Z Cworc ;-.crcducnon Pu; 1. il;e dumnon si .he seernarcgenic c:ic!e and ;?~did:.::~d s p m m s p o n . 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 m d mmagemsni. s a d e m c Press, Xew "_ark
K ~ m c U yJ,
, I, a d I. D. Robsns, i909, Femlitv oi;ayore-ioe, .hybnds.
.:oumd
.
o i i ~ a w n ; i i o gjo: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, m e . md dic:. Cmadian Jounvl
o i Z o o i o ~77: ih4J-56.
Kirchm, A. M., E. M. Grsc. m d E. il Schsustc:. 3M)Oa. Cihm~esm coyote
~ c n v i qp : l t r m 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 ~!oron?)
U
home rmgrs ir. iouthe%rcm C~ioraao.
Canadla Jawnal oiZaoiugy 78:45844.
Zcirn.m. D. i
2 1966 Swnt rnivikng in C m d z e . 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:::
behawor m d m n A g c n e n i .ic~dcierolcPress. New Yak.
K l e m . D. ti..m a .! F. Elsmbcrn. 1973. Carnoanson of c a d ind felid socriii
Hoors:. 3
.
..
.,.
'.
L~unarc. 'V.,ma 3 L. K?Usr 198; Some-imge <lzeaf'zoyoies: .A cxniai
re..?cx. l o m a , ~iWiililifrM m a g e a e x 18:!17-39.
Lawreact. 3. m a 'F 3.3osscn. 1967 Ltultipie c h m s r e r mdysia af C ~ n i s
.'upiir hmns and ,<amilinrs. wch 1 discdsnon i o i me re:atiawiups a i
C ~ n wnyer Amencan iooiog?s;-:133-~1.
L a i ~ e a c c .3 . m d ' N E. 3osser. : 9 6 4 Tllr craniai n-idencz afhybndu3aon
n N e w Engiana C ~ n i rB. i w ~ c r s350:l-13.
Lawreace. 3..md'V 3.F3assen. 1975. Rcist~onshpsoiNilnh Amencan Canrr
3hom-a by imuiupie ciaractervlalysis ci~elemedpopulaoocs.?lges -3-36
.n \.I. W Fox, ed 3 c ,mid i m d s . Vm Y a r m d Rcinhold Xcw Yom.
L e b a Y., A. E i s e h w c r 8. Sanscn. L. D.41r;i. R. 0.?;rrrran, P ; P.
G u m . m d R :C W d y c :YYi.inuogess;on~lfcoyorernrorhondndD*iA
Into jyrnpalmc
. h c r i c m gray w o l ~ o p a i a n o n s 3,oiunon
.
45:lW
:9.
L z h e : . P U. 1 9 7 8 ~Cqore commwcsncn. Pzges : 2 7 4 2 in M.Bckud cd.
Cdyores: Sioiogy, behav:or, md mmngcnent. .Ac=de.mc Press, New Ybik.
L2her. P N. :97% Coyote vacalkiranons: .A lexlcoo m d ;omprisons w h a t h e r
c m d s . .Animal Bzhaviour 26:712-22.
Like:. P. X. 1987. RcpeUenrs m d condirionrd av,xdanca. Plges 5 6 6 1 in 1. S.
Green. ;a. Piote;mg iivesrock born coyotes. L.S. Deamezi of .A.m.
i,dhlre, h:ncuihxu Rtsearch Sc-ce.
US. Shee? Experiment Starroo,
Dubols, ID
L m h x . S. B. 1984a. Manaang coyore h a y prabirns ulth nodethili mchniqurs: Recent aavances in research. Proceedings o f &c Eutcrn Wiidiifc
Damage Canfcrence 1: !05-18.
Linhan. 5. B.1981b. Stroor-lghr mdsirendrilcer for prorec~ngf~nccd-pasnuc
mu m g e rhe-p From Lovote prcdanon. P r a c + c c g s airhe Vertcbmre Pesi
Conference I I:ljiCS6.
L m h a S. B.. m d F. F. ffiowlton. 1967. Detemamng age ofcoyotes hy tooth
c:menrum layen. Journal oiW~laliCe?rlmagement jl:36i 65.
L~nharr,5. R..H. H. B m m m , m d D. 5. 3dsc:. 1968 Fieid c-ialuation o f m
anufemLrj agem. sulberterol. for h b l u n g coyote qruduction. Tiansac tions of rhc 4 o n h . h e n c a n WiiGic Canirrencz 33:316-76.
Lnhan, S. B.. R. T. Sterner, T. C. Cm.gm, and D. R. Henne. 1979. Romandoi guard dogs reduce shcep losses Id C3yOicS: .lprel-ary
cvalmtion.
Joumni o i R m s e Management 3 2 2 3 8 4 1
T.mnan, S. B.. J 3.Robcns. m d ti. !. Dascn. 1982. Elecrric fencmg reducss
coyorrprcdilnon dn paswedsheep. l o r n 1 o f h g c 4 l m g e m c m 2527691
Utah Stare UGvers~ry,Logan.
:(nowiton. i; E. i972. Prellmisrerprctations oiioyore papulanon mcchm~ c s~ 5 t some
h
x m a g m c n r m p i i c ~ o u n s ;ourmi
.
uf Wildlire b i m a ~ e m e n t
36369-82.
.Yl~awltoa, F. F., and E. .LL. Gcse. iY95. Cayate papuiaaon processes re.risired
Pages I 4 in D. Rollins. C. Richarcson, 1B l d e n s h p , K. C a o n . m d S.
H a k e , sds. Cayores ;n illc suarhwerr: -4 compendium of our hiowledge.
Texas Parks m d Wiidiie DepanmenL Austin.
Rnowkon. F F, mdL. C. Stauitvr. 1983. Coyote populanonmrchsolcs: l n c t h e r
look. h g c s 93-1 11 in X. B m c l l , 3. 5. Lsismun. m d 1 LI. ?;rk cds.
Symposium on n ~ i re;nliui~on
d
aiwiidlL5 pupuim>ons. Foissr, Wudiie,
m d R m s c Ex~cnmcnrStahon. Umverritv o i l h h o . >Loscow.
Frede"c:on, Cmada.
i(nowlton, F.F., m d S. L. Wbinrmore. 7'301. M p ccav~ty-toothw d t h mnus h m
.how=,-age a d wiid-~xdghr;oyares deremuned by radiography Wiidlire
Soc~etyYuilcm 29:DY-144.
!(nowiron. F F.. E. &I. Gcse, m d Ll. LM.Jocger 1 9 9 Coyoyare depredaoon ianuoi: .An hteriacc brzveen 5iaiogy md manarmcnr. Journlil a i Range
Lliroaoemcnt .'2:3984!2.
KoeUer. G. >I., m i LL. G. Somocker 1991. Seasonal resource use m o n g
molliazh lionr, iobcz~n.> a dsoyoyares. J o m d ooiMammaicgj 7239;-26.
Kalenosky, 1i.B. l971.iiybndlman bemeen woiimdodcoyacr.Joumaiui>lunm u i a ~ , 532:14619.
Karschgen, L. 1. 1475 k c d 'wbss of coyores in ncrd-cenw >fissaun !Fco<rd
.Ad Projcc: No. W-;3-R27). Llissoun Depamnenr o f Conserranon.
:Qe.+ag, L.W. 1969. ?fir rsr m d fail o i k coyote on lslz Roylilr. >;mmakt
30:24-5:.
Lmprczhr. J. 1978. The reianonshp jewees :bod cornpennon and 'arszmg group m e a sum;
:;sn~vorcs. Z<asck.~?fur T i e r y y c . i o i o ~ r
26:5?7 2:
LLT;ICI.:. 5.. ~ ~ 4 .11:&~... I 9 3 Tie j l i i U I ~iily~ii-s
J SI :
C aC
u ;umon3):
~
,;ommea~. ..oumi af!dmzaaiog~ :A.:ll72--J.
..
L~umlrr.J. tV I98 :. Tcniord .r.ir.;inon m isyoie iccdizadon ntes. : o d o i
Wddhie . m e m e n < -15:767+9.
Linhan S. B., R. i Sterner, G. I Dasch, and J. W Thade. 1984. EBicncy of
l i d r and sound stimuli %ircducmg coyorr ;rreda!ian upon p a s m d sheep.
Prorecr~onEcology b:iS-84.
Linhan. S. B., G.!. Dasch. R. R.!ohnson, J.D. Robens, andC. JP~ckham.1 9 4 2 ~
Elecrromc .h&rhrcliing devices ior reducmg coyote depredation oo domestic
she:?: E E C ~ L under
Y
rmgc conditions m d operationd use. Practrdingr
oithe Vertebnce Pesr Canicicnce 15:386-92.
L m e l l , J. D. C., 1. Odden, LL. E. S m r h R. m r s , and I E. Swenson. i999.
L q c c m v a r e s -bar kill iivesrock: D o ,'problrm indivtduds"re3ily aisc!
Wildliir Suciey Bdicnn 17:698-705.
Llrvaias, J A . 1978. .Llavcmenrs m d hab~tztuse 01' coyotes on the Wichita
Mountams Nanonai Wiidiie ReQugc. Thrsts, Oklahuma Smc Uiuvm~ry,
Stflmrer.
MiiacCacken, J. G. 198!1. Cayore p e y cholce: >\ r q i v Journd ufwtidliic ManIgenenr 53:666-67.
btwCmcken. J. G.. axxi R. hi. E m e n . 1987. Cdyocc feeding suatesies m southmsrern Idaho: O o m a l ioninns
p o o m s u c ;~re&tator?Journal of
. .bv
. m o..
wddliie Mvlanagement 5i:378-85.
> l a h a B. R. 1978. Occumnce u i n c l m s n c c a d s % Nebraska. koccedings
of-be N e b n s l a Academy uiSclence 5 : 1 3 1 3 .
>S~jar,J. T,md 1. .A. Shcrbumr. 1987. IntenpecSc ieiaoansbips of coyotes.
bobcars, and red foxes m wesem M m e . Jaumd ~ i W u d l i f eManagement
51:60&i6.
Mathwg, H. L 1973 Food md popuiation chmcrenstics o i l o w cnjoces. Iowa
State J a u d of Rcscm2-17: 167-89.
McAico, J. K., md D. -4. :(1eoenow. 1978. Prcdanan on a g e i h e q with no
predaior conuai. ; o m d a i X m g e Mmagemmr 3 I: 1 1 1-14~
McBr.de, R. T. 1974. Piedator prorecrion ioilsr for :ivesrack. P ? W n ?io.
3,312,306, :e;.stc:cd
0c:obcr 31, 1974. L S . Zxenr Oficc. WuhmgTon,
-.
UC.
Mc3ndr. R. T 1982. Prebror cunuoi rouc ;;lib. Plrenr Xu. 1,138,386, ;egiscersd Juiv ;7., 1482 1: 5.P3rem Ofice. Wasiunman 3 C .
LIcCjr:::,
3. 1975 Lang-.lirraocz ,;ocaiizanonj .:i;o,8z,rss , Cdno iomns:.
J o m i o i L L u u n d o ~ 56:34>50.
#
L l c C h s . >I. ?. N. 5. 5nnm mu 'V 'V.Siiaw. :9?5 3tr:s ai:oyores = e x h e
h < ; u n w ili SWL-O Uanonal Mcnumcar ma Tucson, . d o n a . South~
~
COYOTE (CUlliS fUbUl1.S)
of nonhem Ycil, Ea,olanC wlth obscnatmns on i m d hybn&. Wildliie
Monogaph 1 7 . 1 4 1
Snow C. I. 1 9 6 7 Silrnc obienarlons on :tic bcha\,lonl m d morp~olog!caldev;lopmeni of soyor; pops . h c r r c m Zoologlrt 7:3ij-i)
soulC,1 . E , D T Bolger.+. C. ilbcns. J. Wnghr,hi. Sounc:, z r d S iiill. !988.
Reconauucrrd dpamlcs of rapia cntincrions o i chaparrcl-iequmog btras
lo urban habtwr ~slandi.Canscrvsnon Biology 2:75-92.
S p ~ ~ g rJ : T~.,1977 lovemrnr patiems o i coyotes ili south-ccnrral W~shingtar.
as d r t e m n e d by rildlo tclcmcny. D~sscnanan,\Vashingmil S h e Uniier
sity, Pullman.
StellHug, J. U , N. L.Gaes, and R. G . Sasses i 9 7 8 Reproductive uhibitars for
coyotc popoi.~tioi~
control: Devcloprncnrs and current ivatus. Pmccedingr
o i r h r Vertcbraa Peit Conference 8:185-89.
Sterner, R r., illd S. A. Shunidkc. I978 Coyote l i a m q e - i a n ~ o lieseaick .X
rcvicw and analysts. Pages 297-325 i n M . Bckorf, ed. Cuyates: Biology,
behavior, and management. icadcmlc Prcss, Yew York.
Thomas, C.U. m d P E. Hughes. 1992. Plague suiveillance by seroloqlcsl testing
of coyotes (Cunis luoiins) in 1.0s Angclrs Coio~ri:Califumia, Joumai of
Wiidlifc Dlssases 28.610--13.
Thoma, N J., W J Forcyt. J. E Evemann, L. A Windberg, arid 1: F.JCnowltun.
1984 S~roprcvalrnccof canine p,uvuvims 111 rvlld coyarcs from Texas,
Llldi, .tud lddi/lo(iY71-1983). Juun~alo f t h e h c r l z a o Vctciltrary.Medica1
.kssocimon 1X5:1233-87.
Thomas, R. E., M. L. Be=& T. J. Quan, L. F. Cirrtci, A. M . Rtirnes, and
C. E. liulila. 1989 Expcnmcntally ii:duced pbamc mfect~un~nthc
rlrirrhem
graa~lloppcrmouse (On),chocny.s Ieuco~us!er)acquircd by consmptlan of
i h c t c d prey lourndl of \Vildlifc Diseases 25477-811.
'Thamto~~,I.
E., I < I<.Licll, ailcibl, J Rc~rdiln.1974. Inrernal parssltcs ofcoyotes
it saiahcrn icnas lounlal of Wiidlifc Disrares 10232-36.
Thurber, J hl , und R. 0. Pe:crson. 1991 Changes io body s h e asbociared with
range enpanilon k [he coyotc (Cunis ioirnni;. Journal of W m ~ a l o g y
72:750--55.
Thurber, J. .VI, R. 0. Pcternio, J. D Woolinoron, m a J. A. Wcctich. ;9'12.
Coyote cocxlstencz w ~ t hvolvcs on rhc Kznai Pcntnsuia, Alaska. Canadian
Journal of Zoology 70:249S~Y8.
T111, I. A , m d 17 1; i(;iowltou. 1983. Eticacy of drnmng in illlcviiiung coy^
uic iieprciiat~onsupon domestis sheep. Jounlal oi Wildlift Lianagemcot
47.Ii)lX-25.
Tinun, R. M , and G. E. Cilmully. 1980. How doyoirs kill shs-p. N,~riuo.ilWool
Grower ;tl:ILl5.
Timm. R. Ll , and K. 11. Schtmdt. 1989 blanaeelncat oroblcrns cncounrered
(I. . ..
. ! :
! ' , I
. J.' '-.,:.,,:. :I,,..-:.,!
!.
..!.,,.,,.
',5L,:.
, a ? : .,..>, 8 . ,A...,.A:.
,.,,,I.,<C
c,,.
.A~<*..l.,"
>:r. .< ,.<.::::.
I..,.,,...
<;,-r. .LA: : - ' ('5 l>::brr,::.r
Agnculturc.
Todd A. W 1985 Dernagraphlc w d dicory coinparisons offarest and farmiand
coyote, C;lnis lununs. popularions m Aibibcna. Canadian Fir!d-Naruraiist
99:163-73.
Todd A. W., and L. U Keltll. 1983. Cilyotc demography dunng a snowshoe harc
decline in Albma. Jawnal of Wildlife bliinagrment 47.39W0.1.
Todd A . W, L. B.Kclrh, andC. A. Fischci 1981.Populat~onccolagyofcoyotes
dunng d liuctuiit~unof snowshoe harcs.Journal of WililliR Management
45:02910.
Trainer, D. 0 , and F. F Knoivlton. 1968 Scrolog~cc v ~ d ~ n ofdiszases
ce
mTexnr
coyolcs. Journal u t Wildliie Miinagcnlcnr 32:961-83.
Tiilkowsh, W bl. 1980. Monnliry patterns ofiadlo-marked coyores in Tacksou
liolc, Wyommg. Disscrmon, Univrrslty of Llaasacbu~ctts..Anhcrsr~
U.S. Dcpamrrenr of.k&?lculTWc. 1994. Final cnvtronmmrd impact sratemzn!.
Animal Damage Control Prognm, Val., 2. h u m s 1 and P i a t Hcalm Inspection Scrvlur, Washlogton, DC.
U S . Depamocnrafh&mculrure.1995 Shcepandgoiirprrdarurloss. US. Dcpmmen1 u i Agnculturr, National Agncuitunl Stausrics Boar4 Vf&tlmgran,
.
,
,
.,
.,
--
"C.
U S . Fish and W~ldlifcScrv~ce.1978. Predator ailmagc in the west: .4 study
o f coyote milnagcrilrrii altrrnativcs. I;S. Fish md Wildlife S m i c e ,
Washington, DC.
VarlVurm, D , m d S. E. Ilompson, Jr. 1982. Oppoirunlsnc feeding by coyotes.
Nanhwrsrrrn Science 56: 131-35.
Van Wormer, I.1964 Thc world ofthr coyore. Ljppincurt, Philadelphka.
Voig. U. K.,and B. D E a l i . 1983 Avotdancr of coyotes by red fox funilics.
J o u r n ~ of
l Wlldbfe Liauagcmcnt 47:852-57
Wadc. D. 3. 1978. Coyore damage: . survey o i its name and scope, conool
measures and :heir dppiicntion. a g e s 3 4 7 4 8 m M. BekaJT. sd. Coyoies:
Biology behav~arm d manaycmcnr Academic Press, 4 e w York.
Wagner, r H. 1988. Predator connol aod the she;? iuidusny, Rrglua Book,
Ciaremonr, CA.
481
Wagner. K. K., andhl. R. Sonaver 1999 Etiect ofprevenvarivecoyote hunringon
sheep :osari ro covore piedarlon. J o w i l oi\Vildhic Clunsgcmenr 63 606..
12.
Walsh, P. B., and J. LL Inglis. 1989. Seasonal and ,lie1 rare a i rpantanzow
vocnlizar~onin cavotcs m soudi Texas, Jownal of \l-aiorv
70:109-71.
Ysw ~ m n b v i c kFrcdencron, Canada.
Wewer, J. L. 1977. Coyore-food base rebarionships m lackron Bole: Wyoming.
Thesis, Utah State Uwersiry, Logan.
Weaver 1 L. 1979 Influrace of sk c m o n upon cayotc populahoor in Jackson
Hole, Wyornmg. Pages 152-57 m hi S.Buycr and L. D. Hayden-Wing,
cds. North irnencm c k Ecalugy. behavior and mmagemcnt Gmversi~y
of Wyomicg, Lanmie.
Wcwci, J. L., 2nd S. W Hilfman. 1979. Dlffrrenrrill Jerecrabilin, ofrodmts in
gray wolf icvts. Jounnl of Wildlife &lunagcrucnt 43:783-86.
Wzils, M.C. 1978. Coyote s m s c s in predarion: Envsronmentai influences on
then relanve use. Behavlarvl Praccedings 3.149-58.
Wells, M. C., and P. N,L;:mcr. 1978. l h e ieiatlvc unpanaoce o i m e distance
senses in ioyotc predarory b c h a v ~ ~ uh m
r l i l i Brhavtour ?b:25i-58.
Wells, >[. C., and M. BrkaR 1481. .An obiervaiionill smdy of s c z n t m a r h g by
wild coyotes. Animal Behavlour 9:332-50.
Weils, .M. C., and M. BckolP 1982. i'rshc~onby wlld coyotes: Bchavsoral and
ecological analyses. Journal ofblazmaiogy 63:118-27.
Wlntr, P J., K. Ralls, and C A. Vandexbtlt Whirc. 1995 Overlap in habitat
and food use between coyotes and Sm J i a a q h kit foxes. Saurlrwsstem
tsi ~
l)3
4
7
1
)4
0
:tsi K
a
ln
T
u
r
4!1:34249.
Williams, E S., E.'r. Thornc, M, I G. Appci, ~ n D.
d W Bclilbky. 1988. Canine
disrentpei in black-footed <errcia (IWusrei~~
nigrrpr~lficm Wyoming. Journal o i Wildlife Diseoses 24:385-98.
Wilson,P. J., S. Grewal, I. D.Lawfbrd I N. M.Hrai, A. G . Gmnaciil, D.Penaock
J. B Thebcrgc, >I. :
Theberge, D. R. big!, W. Waddell, R. E. Chambers,
P C. Payuct. G. Gotile:; D. f l u s , and B. N.Wllire. 2000. DNA profiics of
h c eaitcrn Cmadian wolf and ihc red waifprovide evidence for a comnlon
evoliltionwy lusrory mdcpende:~t of the g a y woll Cmadlm Joilrnal of
Zoology 78:215646.
Wmdbcrg, L. h 1995. Demugiapily of a hlgb-density coyotepopuiation. Canadian lour.aal aiZoolagy 73942-54.
Widberg, L. A. 1996. Coyote respanscr to v~suui*rid olfactory stimuli related
:o iarmliuianlv w ~ t han m a Canadian loumnl afZoolary
. :I:?L4&54~
Wimlbrrg, L A , and F. F. Knou~lton.1988. blaiiagemcnr implications of'coyore ipacmg pinems m sourhwn Texas. Journal of WiidliCe bfanugcmmr
52:632-10.
Windberg, L. .A, and 1: F Knowitan. 1990. Relative vulnerab~lityof coyotes to
some capture procrdilres. Wildlife Suc~cryBulletin 18282-90.
W-~ndburg,L. A , ard C. D .\litchsli. !990 \Viaicr diets afcoyotcs in rclanan
to prey abundmcc in southern Texas. Journal of >iarnmalogy 71:43917.
Windberg. I.. A , M.L. Anderson, a n d R M. Engemim. 1985. S w l v u l ofsoyares
in southcm~exas.Journal of Wildlife Managemncnt 19:301-7.
Wmndberg, L. 4 , 11. M. E n g e m s , and J. F. Broniaglnn. 1991 Body size and
condit~or~ofc~~ynres
msuuthern ];eras. JournalofWildlifiDisenres27:4752.
Windberg, L. A,, S. M. Ebbcrt, and 8. Z Kelly 19978. Population charactenstics of coyotes (i'zi,iir lornzns) in the nortltern Cliil~u.lhuandrscn oiNew
)rtexico. .lmencm hlidland Yaruralist 138:197-207.
Whdbcrg, L. A , i F Knawlton. S. >I. Ebben, a n d B . 1 Kelly. 1997b. .kspecrs or'
coyors predation an Angora goars. Journal of Range Management 50:226..
JU.
Winsom, M.J.. S. P Green, R. M Cowin, a n d E K.Frirrcll. 1991.DiroJiiario
~nanitlsin coyotes and foxes in \.lissaun. Journal of Wtldlife Diseaes
?7:16649.
Woodruff, R. A,, and B. I..Kellcr. 1982. Dispersal, daily activity, and hooine
n n g e of coyotes in routhessrern Idaho. Yonhwest Science 56:199-207.
Wmter, D. H., and K Bciurschkr, K. I 9 6 8 Comparative cymgenetic studies in
the ordcr C h o r a . Chramosuma ?4:336~82.
Wimagernent Insnrute, Washington. DC
Ltrnc BEKOFT,
Depaiment crf Ei~vironmnru!. Popuiarian, and Organismic
Biology, Umvenity of Colorado, Boulder, Coiurada 80309-0334. Emall: marc.
brkoif@colurado cdu.
EMC hl. GLSE,Nalional Wildlife Research Ccnter, Department of Forest,
Range, and Wildlife Services, Utah Sraie Umvcrslty, I.ugal, Utah 84322-5295.
Emad: cgesc@cc.usu.edu.