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Social organization of the coyote in relation to prey size Bowen, William Donald 1978

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SOCIAL ORGANIZATION OF THE COYOTE IN RELATION TO PREY SIZE by WILLIAM DONALD BOWEN B.Sc, University of Guelph, 1971 M.Sc, Univ e r s i t y of Guelph, 1973 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA August, 1978 William Donald Bowen, 1978 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f a n a d v a n c e d d e g r e e a t t h e U n i v e r s i t y o f B r i t i s h C o l u m b i a , I a g r e e t h a t h e L i b r a r y s h a l l m a k e i t f r e e l y a v a i l a b l e f o r r e f e r e n c e a n d s t u d y . I f u r t h e r a g r e e t h a t ' p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s m a y b e g r a n t e d b y t h e H e a d o f my D e p a r t m e n t o r by h i s r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t b e a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . a r t m e n t o f 2~Oolc T h e U n i v e r s i t y o f B r i t i s h C o l u m b i a 2075 Wesbrook Place Vancouver, Canada V6T 1W5 D a t e 3o kk/ - 0 7 2 i i ABSTRACT The t y p i c a l s o c i a l unit i n coyotes, Canis l a t r a n s , i s the adult heterosexual p a i r ; young usually disperse during t h e i r f i r s t year. In some populations, however, some young delay d i s p e r s a l u n t i l a f t e r the b i r t h of a subsequent l i t t e r . In addition to t h i s v a r i a t i o n i n s o c i a l i t y , numerous studies have indicated marked v a r i a b i l i t y i n the proportion of ungulates i n the coyote d i e t . The e c o l o g i c a l correlates of v a r i a t i o n i n coyote s o c i a l structure were previously unstudied. Thus, I used the coyote as a test of the hypothesis that s o c i a l i t y i n large mammalian carnivores i s an adaptation allowing the e x p l o i t a t i o n of large prey. I studied the s o c i a l structure and foraging ecology of coyotes i n Jasper National Park, Alberta, between 1974 and 1977. Forty-three coyotes consisting of 10 adult males, 10 adult females, s i x j u v e n i l e males and 16 j u v e n i l e females were captured 51 times. Twenty-six coyotes (>_ age 6 mon) were equipped with r a d i o - l o c a t i o n transmitters to provide information on s o c i a l structure and systems of land tenure. A large majority of the coyotes i n the population were marked or could be otherwise i d e n t i f i e d . To determine coyote d i e t s , I c o l l e c t e d 1,967 known-aged feces from through-out the year. In winter, the Jasper population consisted of approximately 59 percent packs, 17 percent residents p a i r s , 10 percent s o l i t a r y residents and 15 percent transients. Packs consisted of three to eight adults, yearlings and non-dependent young. Radio-telemetry,data and observations showed that pack members frequently t r a v e l l e d , s l e p t , foraged together and cooperated i n t e r r i t o r i a l defense. Agonistic i n t e r a c t i o n s between pack members i i i r e f l e c t e d t h e i r s o c i a l status within a dominance hierarchy. In coyote packs, I found a clear d i v i s i o n of labour i n the i n i t i a t i o n of t e r r i t o r i a l defense, scent marking and co n t r o l of group t r a v e l . Packs and pairs defended well defined t e r r i t o r i e s of approximately 2 8 to 20 km i n winter and probably throughout the year, whereas s o l i t a r y residents occupied.home ranges which they did not defend. Males and females scent marked t h e i r t e r r i t o r y throughout the year. The density of scent marks at the edge of the t e r r i t o r y was approximately twice that i n the center. This was accomplished by reducing the distance between scent-mark locations and increasing the percentage of multiple marks. The scent marks of neighbours were not avoided, but vigorously marked. I found circumstantial evidence that coyotes respected t e r r i t o r i a l boundaries demarcated by scent marks. In winter, coyote diets consisted p r i m a r i l y of ungulates (67 percent), small rodents (23 percent) and other mammals (7 percent). Young cervids and adult ungulates comprised about 50 percent of the d i e t i n summer. Columbian ground s q u i r r e l s and small rodents were the other p r i n c i p a l foods. Judged against an average of 15 other studies, the mean s i z e of prey eaten by coyotes was larger i n Jasper than elsewhere. In Jasper, the percentage of mule deer i n the winter d i e t varied d i r e c t l y with coyote pack s i z e and mule deer density. However, pack s i z e accounted f o r more of the v a r i a t i o n i n the amount of mule deer i n the d i e t than mule deer density. In contrast, the percentage of elk i n the winter di e t was independent of pack s i z e . Since elk were scavenged, t h e i r occurrence i n the winter d i e t was p r i m a r i l y a function of the number dying within each coyote t e r r i t o r y . I suggest that the success of coyotes i n i v k i l l i n g mule deer::was increased by group foraging. Another possible advantage of cooperative foraging was an.increased a b i l i t y to defend an ungulate carcass against conspecific competitors. Packs had greater access to food and fed longer than s i n g l e coyotes. There i s a c o r r e l a t i o n between group l i v i n g i n coyotes and the propor-t i o n of large prey i n the d i e t . A l t e r n a t i v e hypotheses that could account for the v a r i a t i o n i n coyote s o c i a l structure i n d i f f e r e n t environments are examined. I conclude that increased s o c i a l i t y i n coyotes i s an adaptation allowing more e f f i c i e n t capture and/or defense of ungulate prey. V TABLE OF CONTENTS Page ABSTRACT i i TABLE OF CONTENTS v LIST OF TABLES v i i i LIST OF FIGURES x i ACKNOWLEDGMENTS x v i 1 INTRODUCTION 1 2 STUDY AREA 5 Geology and S o i l s 5 Climate 5 Vegetation 8 Fauna 10 History of the Coyote Population 11 3 GENERAL METHODS 18 4. CAPTURE DATA AND TELEMETRY 26 Capture Data 26 Telemetry Studies 26 5 POPULATION BIOLOGY 30 Numbers 30 Reproduction 33 Mo r t a l i t y 38 6 HOME RANGE AND PATTERNS OF SPACE USE 46 Methods 46 Results , 47 v i Home range as the asymptote of the observation-area curve .. ... 47 Home range f i d e l i t y . 55 Dispersal 56 Size of home range. 58 D i s t r i b u t i o n of coyote home ranges..... . 61 Discussion 79 7 SOCIAL ORGANIZATION 85 Results 85 A s o c i a l c l a s s i f i c a t i o n of coyotes i n the population 85 Transients 87 S o l i t a r y residents 92 Resident pairs 96 Packs 96 Composition and s t a b i l i t y of coyote packs 104 Athabasca Pack 104 Maligne Pack 106 Dominance r e l a t i o n s h i p s within packs 109 D i v i s i o n of labour 113 Genetic relatedness of pack members 115 Land tenure 118 Discussion 121 v i i 8 SCENT MARKING 127 Methods 128 Results 128 Types and frequency of occurrence of scent marks. 128 D i s t r i b u t i o n of scent marks " 130 Temporal d i s t r i b u t i o n of marking... 136 Stimuli e l i c i t i n g scent marking 140 Response to scent marks of neighbours. 146 Discussion 147 9 DIETS, FEEDING AND FORAGING BEHAVIOUR. 151 Methods 151 Results 153 Diet 153 Resident p a i r and pack d i e t s . 162 Comparison: of adult and pup die t s 177 Feeding behaviour at ungulate carcasses.... 179 Ungulate predation 185 Discussion 190 10 SOCIAL ORGANIZATION IN RELATION TO ECOLOGY 196 LITERATURE CITED 211 APPENDIX I 225 APPENDIX II • 228 APPENDIX I I I • 229 v i i i LIST OF TABLES Table Page I Summary of c l i m a t i c information recorded at Jasper townsite f o r the period, 1973-1976. 9 II Summary of vehicular t r a f f i c i n Jasper National Park during the period, 1960-1971 17 II I Results of capture program i n Jasper National Park, 1974^-1976. Number of coyotes radio tagged i s given i n parentheses 27 IV Summary of radio-telemetry data for 26 coyotes captured i n Jasper National Park 29 V Summary of coyote l i t t e r s born on the study area from 1974 to 1976 36 VI Number of coyote m o r t a l i t i e s i n the Athabasca River Va l l e y during the period, 1974-1977 39 VII Number of radio locations and days required to reach an asymptotic index of area u t i l i z e d f o r 26 coyotes i n Jasper, Alberta 50 VIII Time of d i s p e r s a l and distance t r a v e l l e d by 12 coyotes of various ages 57 IX Home range s i z e of male and female coyotes i n Jasper National Park 59 X C l a s s i f i c a t i o n of coyotes based on s i t e f i d e l i t y , group s i z e , group composition, and gregariousness 86 i x Table XI Home range length of males and females of four resident pairs based on observations 97 XII The percentage of coyotes observed i n groups of d i f f e r e n t sizes i n summer and winter, 1974-1977 100 XIII The percentage of observations of coyotes i n groups of d i f f e r e n t sizes i n summer and winter, 1974-1977 101 XIV A summary of the s i z e and composition of the Athabasca Pack from July, 1974 to February, 1977 105 XV A summary of the s i z e and composition of the Maligne Pack from July, 1974 to February, 1977 107 XVI Summary of snow-tracking data from f i v e coyote packs and two mated pairs i n Jasper National Park (scent marks within 20 m of k i l l or carcass excluded) 131 XVII Comparison- of rate of scent marking at the edge and center of a t e r r i t o r y 133 XVIII Number of days each month that nine, adult females were observed scent-marking 139 XIX C l a s s i f i c a t i o n of coyote foods by s i z e (weight). Average weight of each species given i n parentheses 154 XX Percent frequency of occurrence of major items i n 1,967 coyote feces from 1974 to 1977 155 XXI Percent frequency of occurrence of d i f f e r e n t s i z e prey i n 1,463 adult coyote feces 161 X Table XXII A comparison of the percent occurrence of three prey species i n Maligne Pack feces to a sample of feces c o l l e c t e d i n the pack's t e r r i t o r y (of unknown or i g i n ) 163 XXIII Percent frequency of occurrence of each food cl a s s i n the winter feces of coyote pa i r s and packs 164 XXIV Percent frequency of occurrence of each food class i n the summer feces of p a i r s and packs... 165 XXV Number of vehicular ungulate deaths per coyote-group t e r r i t o r y i n winter, 1974-75 and 1975-76 combined 167 XXVI Comparison of the proportion of ungulates i n the winter d i e t of three coyote packs between 1974-75 and 1975-76. Frequency of occurrence i n feces given i n parentheses. 175 XXVII Comparison of the r e l a t i v e frequency of each food-s i z e class i n adult and pup feces c o l l e c t e d from May to July, 1974-1976 178. XXVIII The number of v i s i t s i n which non-satiated residents and v i s i t o r s fed upon ungulate carcasses i n d i f f e r e n t s o c i a l contexts 181 XXIX Percent frequency of food classes 1 to 4 (see Table XIX) i n coyote diets based on 18 studies 205 x i LIST OF FIGURES Figure Page 1 Map showing the l o c a t i o n of the study area i n Jasper National Park, A l b e r t a 6 2 Approximate winter d i s t r i b u t i o n of Ovis canadensis i n the Athabasca River V a l l e y , Jasper National Park 12 3 Number of coyotes k i l l e d i n Jasper National Park during periods of predation control ( • ) and by motor vehicles ( O ) from 1934 to 1976 15 4 A l l subadults and adult coyotes captured were equipped with a radio transmitter i n the form of a c o l l a r and were given an ear tag and neck tag . 21 5 Drawings of coyote pelage c h a r a c t e r i s t i c s used to i d e n t i f y c e r t a i n members of the population 24 6 Minimum number of coyotes a l i v e i n July and March, 1974-1976, i n the Athabasca River V a l l e y . , . Data a v a i l a b l e f or only f i v e of nine coyote groups. ^The l i t t e r of three groups was not found, however, I assumed two pups a l i v e per group at the time of the census 31 7 Percentage of t o t a l sightings which were of known coyotes as a function of months during the winter of 1974-75 ( O ) and 1975-76 ( • ). Number associated with each symbol represents the t o t a l number of sightings per month 34 x i i Figure Page 8 Pattern of h a i r loss which i s c h a r a c t e r i s t i c of advanced scarcoptic mange i n f e c t i o n 42 9 Relationship between an index of area u t i l i z e d and time f o r adult, female C3 from October, 1974 to July, 1975 51 10 Relationship between an index of area u t i l i z e d and time f o r adult, female C4 from October, 1974 to July, 1976 .. .. 53 11 Relationship between home range size, and pack s i z e during the winters, 1974-75 ( • ) and 1975-76 CO). a. Home range s i z e estimated from the d i s t r i b u t i o n of observations.... ^Home range s i z e corrected f or water area 62 12 The s p a t i a l d i s t r i b u t i o n of resident coyotes on the study area i n the winter of 1974-75. See text for explanation of map 64 13 The s p a t i a l d i s t r i b u t i o n of resident coyotes and active dens ( • ) on the study area i n the summer of 1975 66 14 The s p a t i a l d i s t r i b u t i o n of resident .coyotes on the study area i n the winter of 1975-76 68 15 The s p a t i a l d i s t r i b u t i o n of resident coyotes and ac t i v e dens ( • ) on the study area i n the summer of 1976 70 x i i i Figure Page 16 D i s t r i b u t i o n s of radio l o c a t i o n s f o r adult females C3 ( • ) and C6 ( A ) during.the winter, 1974-75. The number above each symbol indicates the number of times each coyote was at various locations 73 17 Frequency d i s t r i b u t i o n s of radio locations of females C4 and C l l during the period"October, 1975 to July, 1976. Data f o r each animal are plotted on the same coordinate space. - 77 18 The number of coyotes i n each of four s o c i a l classes i n March, 1974-75 (open bars) and 1975-76 (shaded bars). Percent of t o t a l number of animals i s given above each bar. 88 19 Observation-area curve of subadult female C22 from November, 1975 to March, 1976. Every second radio l o c a t i o n i s p l o t t e d 90 20 Observation-area curve of adult male C14 from A p r i l , 1975, to May, 1975. Every second radio l o c a t i o n i s p lotted 93 21 Frequency d i s t r i b u t i o n of radio locations of s i x members of the Athabasca Pack from A p r i l to September, 1975. Data f o r each animal are.-.plotted on the same coordinate space 102 x i v Figure Page 22 Dominance r e l a t i o n s h i p s among members of the Athabasca Pack i n winters 1974-75 (A) and 1975-76 (B) and the Maligne Pack i n winter 1975-76 (C). An arrow indicates that one coyote dominated another, whereas a dotted l i n e indicates that animals were of equal rank 110 23 Mean rate of scent marking as a function of group s i z e at the edge (open bars) and i n the center (shaded bars) of a t e r r i t o r y . Sample s i z e and S.E. are indicated above each bar 134 24 Expected mean number of scent marks per km ( O ) assuming a l l coyotes mark at the same rate and mean observed rates of marking ( • ) as a function of group s i z e 137 25 Relationship between number of scent marks per coyote-km and months, of the year at the edge (•) and i n the center ( O ) of the t e r r i t o r y . Sample s i z e indicated above each symbol 141 26 V i s i t a t i o n of scent marks by the Maligne Pack along a .5 km section of road near the edge of t h e i r t e r r i t o r y . Sites marked ( • ) and s i t e s v i s i t e d but not marked ( O ) are indicated. Absence of a symbol means that a scent mark was neither marked nor v i s i t e d that day 144 XV Figure Page 27 Seasonal v a r i a t i o n i n the percent frequency of Columbian ground s q u i r r e l s ( A ) , mice ( • ), young cervids ( A ) , and other ungulates ( a l l . species >6 mon of age; O ) i n coyote feces 157 28 Relationship between the percentage of ungulates i n winter feces and pack s i z e during 1974-75 ( • ) and 1975-76 ( O ). Data from s o l i t a r y residents ( • ) combined f o r the two years 169 29 Relationship between the percentage of elk i n winter feces and pack s i z e during 1974-75 ( • ) and 1975-76 ( O ) 171 30 Relationship between the percentage of mule deer i n winter feces and pack s i z e during 1974-75 ( • ) and 1975-76 ( O ). 173 31 Mean time spent feeding at ungulate carcasses by residents (A) and v i s i t o r s (B). Ninety-five percent confidence l i m i t s and sample s i z e are given above each mean...PA: Pack alone at carcass, SRA: Single r e s i -dent alone at carcass, P-V: Pack and v i s i t o r ( s ) present, SR-V: Single resident and v i s i t o r ( s ) , SVA: Single v i s i t o r alone, V-V: Several v i s i t o r s present, SV-SR: Single v i s i t o r and si n g l e resident present 183 32 Wounds sustained by an adult female mule deer during a coyote predation attempt 187 x v i ACKNOWLEDGMENTS Many people contributed to t h i s study, and I regret that i t i s impossible to acknowledge everyone who helped i n one way or another. Dr. Ian McTaggart Cowan encouraged me to attend The Un i v e r s i t y of B r i t i s h Columbia and introduced me to the challenging study of large vertebrate predators. 1 gained much from our weekly meetings during the 10 months p r i o r to i n i t i a t i n g the f i e l d work. During my stay i n Jasper, h i s v i s i t s were the source of needed encouragement to an often f r u s t r a t e d young ec o l o g i s t . I am deeply indebted to him f o r h i s enthusiastic support. This study would not have been possible without the support of Parks Canada, The Department of Indian and Northern A f f a i r s . Jasper National Park Superintendent Rory Flanagan and Assistant Superintendent Bruce Wilson ass i s t e d greatly i n providing Park Annual Reports and access to various park f a c i l i t i e s . I am most g r a t e f u l f or the cooperation and support given by Bob Haney, Head of Resource Management Function, Warden Service. Many other wardens also made valuable contributions to the study, but the help of several of these deserves s p e c i a l mention. Duayne Martin showed an early i n t e r e s t i n the study and a s s i s t e d i n many ways throughout the research. A l f i e Biirstrom was a constant source of information, p a r t i c u l a r l y regarding the l o c a t i o n of coyote dens. Marvin M i l l e r and Robert Watt gave f r e e l y of t h e i r time arid knowledge of the area. I w i l l long remember the friendship and help extended by each of these men., I am also indebted to Roy Routledge f o r h i s prompt and s k i l l f u l r e p a i r and maintenance of the radio-telemetry equipment. Jack P l e c k i t i s and Doug Stewart were able and enthusiastic f i e l d x v i i " a ssistants i n 1974-75 and 1975-76 r e s p e c t i v e l y . Many of the observations reported i n the thesis were made by them. In p a r t i c u l a r , Doug Stewart c o l l e c t e d much of the scent-marking data on the Maligne Pack. I am g r a t e f u l to Daphne Hards f o r preparing t h i n sections of coyote teeth; Nasar Din, C u r a t o r i a l Assistant of the Cowan Vertebrate Museum, The University of B r i t i s h Columbia, for the use of museum specimens and i d e n t i f i c a t i o n of h a i r ; Darlene Belford f o r help i n computer program-ming; and L e s l i e Borleske for kin d l y typing the t h e s i s . Drs. I. McT. Cowan, H. D. Fisher, H. C. Nordan, J . N. M. Smith, and C. J . Walters kin d l y reviewed the manuscript. I am g r a t e f u l f o r t h e i r comments and suggestions. The thesis i s much improved as a r e s u l t . F i n a l l y , and most e s p e c i a l l y , I thank my wife, E l l y , f o r her i n t e r -est i n the research and for her support throughout the study and p a r t i c u -l a r l y i n the f i n a l stages of w r i t i n g . 1 1 INTRODUCTION The s u r v i v a l of an animal depends c r i t i c a l l y upon i t s a b i l i t y to f i n d food and to ex p l o i t i t adequately. It follows then that the s o c i a l organization of a large mammalian predator with few natural enemies may be p r i m a r i l y an adaptation to i t s food. Adaptations to deal with predation and other factors should be superimposed upon t h i s primary organization (Crook 1964). Group-living carnivores have evolved independently several times, and at l e a s t 16 of 253 extant species e x h i b i t a high degree of s o c i a l i t y (Wilson 1975). Nevertheless, highly s o c i a l carnivores are uncommon. Indeed i t i s s u r p r i s i n g that they should e x i s t at a l l , given that there i s no automatic or u n i v e r s a l benefit to be derived from group l i v i n g (Alexander 1974). On the contrary, there appear to be widespread d e t r i -ments associated with increased s o c i a l i t y , namely 1) increased:'.intensity of competition for resources, including mates, 2) increased l i k e l i h o o d of disease and parasite transmission, and 3) increased conspicuousness e i t h e r making a species a l e s s e f f e c t i v e predator or more vulnerable as a prey (Alexander 1974). Thus, we must explain the evolution of s o c i a l groups i n the face of the above opposing s e l e c t i o n pressures. Alexander (1974) stated that the s e l e c t i v e background of group l i v i n g should involve 1) reduced s u s c e p t i b i l i t y to predation, 2) increased e f f i c i e n c y of e x p l o i t i n g a needed resource, or 3) extreme l o c a l i z a t i o n of a resource. Alexander went on to suggest that group l i v i n g evolves because one or some combination of the above s e l e c t i v e forces enhances -2 . ( the f i t n e s s of in d i v i d u a l s accepting the detriments of group l i v i n g above the f i t n e s s of s o l i t a r y i n d i v i d u a l s . Bouliere (1963) observed that most predators se l e c t prey about t h e i r own body weight or less and have a s o l i t a r y mode of l i f e . However, a / small group of highly s o c i a l predators r e g u l a r l y take prey species often exceeding several times the weight of an i n d i v i d u a l predator. The three most l i k e l y hypotheses to explain group foraging are: 1) the advantage of group foraging may be to ex p l o i t a food resource unavailable to a single predator ei t h e r due to the large s i z e and/or speed of the prey (Kruuk 1972); 2) group foraging i s favoured when o resources are scattered, or concentrated food i s located too infrequently by single animals (Alexander 1974) ; and 3) group foraging i s favoured when there i s competition over large economically defendable food items such as ungulate carcasses (Eaton 1976, Schoener 1971). There i s general agreement that within the Canidae most species (e.g. Vulpes vulpes) studied thus f a r are rather s o l i t a r y . Conspecifics s o c i a l i z e p r i m a r i l y during the breeding season, have rather uncomplicated but highly stereotyped v i s u a l displays (Kleiman 1967, Fox 1970), do not possess behaviours for the maintenance of cons p e c i f i c groups and thus cannot exp l o i t prey species whose body weight i s much greater than that of t h e i r own. Consequently, the d i e t consists p r i n c i p a l l y of small items such as mice, lagomorphs, and f r u i t s . In contrast to t h i s , the highly s o c i a l wolf (Canis lupus) possesses behaviour patterns which from early i n l i f e (Fox 1972, Bekoff 1974) enhance pack formation and 3 maintenance and allow the t r a n s i t i o n from a s o l i t a r y to a s o c i a l e x i s t r ence as dictated by changes i n food supply or prey type (Kruuk 1972). In most areas, group predation of large ungulates i s the r u l e f or t h i s species (Mech 1970). Thus, the red fox and wolf represent opposite extremes of a phylogenetic continuum.' of s o c i a l i t y i n which food s i z e r e l a t i v e to the si z e of the predator may play a major r o l e . Available information, although meagre, suggests that the coyote, Canis la t r a n s, i s intermediate between the red fox and wolf i n s o c i a l behaviour (Kleiman 1967; Bekoff 1972, 1974; Fox 1975), s o c i a l organiza-t i o n (Fox 1975) and foraging ecology (Gier 1975). Fox (1970) showed that the f a c i a l expression of the wolf and coyote are much more v a r i a b l e and exhibit greater degrees of graduation and simultaneous combination i n contrast to the more stereotyped expressions of the foxes, Vulpes, Urocyon, and Alopex. Fox's data demonstrate c l e a r l y that -the coyote and wolf form one behaviour group while the foxes form another. Studies on canid s o c i a l i z a t i o n and play behaviour further i l l u s t r a t e differences i n the ethology of coyotes, wolves and foxes (Fox 1970, Bekoff 1974). Thus, i t appears that coyotes possess the behaviour permitting a su b s t a n t i a l degree, of cooperative foraging behaviour. I f ' t h i s i s true I would expect to f i n d that coyotes should be able to adjust t h e i r s o c i a l organization to changes i n the s i z e of a v a i l a b l e food. Further, I predict that the degree of s o c i a l i t y i n coyote populations under natural conditions i s p o s i t i v e l y correlated with the average /size of prey eaten. 4 This study addresses the following questions: 1) How are coyotes organized i n time and space? 2) What i s the nature of the food eaten by coyotes? and 3) How do coyote group s i z e and s o c i a l structure vary with the s i z e of food eaten? These questions provide a means of t e s t i n g hypotheses concerning the r o l e of cooperative foraging i n the evolution of canid s o c i a l i t y . I selected Jasper National Park as the study area for two reasons. F i r s t , the park supports a high ungulate biomass. Thus Cowan (1943) and Hatter (1945) reported that the r e l a t i v e frequencies of ungulates i n coyote winter feces were 70.0 and 45.4 percent r e s p e c t i v e l y . These percentages are among the highest reported f o r any coyote population thus far studied (see Weaver 1977). Second, coyotes i n the park have not been co n t r o l l e d for approximately 25 years. Thus, both conditions considered necessary and s u f f i c i e n t f o r the expression of cooperative coyote groups were s a t i s f i e d i n Jasper National Park. 5 2 STUDY AREA Delimited to the west by the Continental Divide, Jasper National Park, Alberta (6720 km2, 53° north l a t i t u d e ) l i e s e n t i r e l y within the Front and Main Ranges of the Rocky Mountains. The main study area was 2 120 km i n the Athabasca River V a l l e y between Jasper townsite and Pocahontas Warden Station (Fig. 1). Less intensive work was conducted i n the Maligne and Miette River Valleys and south of Jasper to Buffalo 2 P r a i r i e ; a t o t a l area of approximately 300 km . Geology and S o i l s The Front and Main Ranges are complex, folded sheets of Paleozoic carbonates and shales and Mesozoic shales, sandstone and limestone (McKay 1952). Rising abruptly to jagged peaks of 3000 m, the mountains tower above the v a l l e y s below (975 m) creating one of the most spectacular landscapes i n North America. The v a l l e y s are covered with blankets of g l a c i a l t i l l s , g l a c i o f l u v i a l and p o s t g l a c i a l alluvium deposits reworked by wind and water act i o n . These s o i l s are x e r i c , undeveloped and nutrient-poor i n the grassland areas of the Athabasca Valley (Stringer 1969, quoted i n Stelfox 1974). Brisk, f o e l n winds cause marked snow abl a t i o n i n winter and have produced extensive loess deposits i n the v a l l e y (Stringer and La Roi 1970). Climate The Athabasca Valley has warm summers, a long snow-free period and 6 F i g . 1. Map showing the l o c a t i o n of the study area i n Jasper N a t i o n a l P a r k , A l b e r t a . 7 8' c o l d , r i g o r o u s w i n t e r s owing t o t h e a l t e r n a t i o n of v e r y c o l d A r c t i c systems and warm P a c i f i c a i r masses. Weather d a t a f o r J a s p e r t o w n s i t e a r e summarized f o r t h e y e a r s 1973-1976 i n T a b l e I . J u l y i s t h e warmest month and J a n u a r y t h e c o l d e s t w i t h extreme low t e m p e r a t u r e s o f -40°C. On a v e r a g e t h e months o f June, J u l y , and August a r e f r o s t - f r e e . P r e c i p i t a t i o n v a r i e s c o n s i d e r a b l y i n d i f f e r e n t a r e a s o f t h e p a r k . The A t h a b a s c a V a l l e y , as i n d i c a t e d by J a s p e r t o w n s i t e d a t a , i s t h e d r i e s t , r e c e i v i n g o n l y 34.9 cm p e r y e a r . By c o n t r a s t , W i l l o w Creek i n t h e n o r t h w e s t s e c t i o n o f t h e p a r k r e c e i v e s a p p r o x i m a t e l y 73 cm o f p r e c i p i t a t i o n p e r y e a r ( L a n d a l s and K n a p i k 1972, C. W. S. r e p o r t ; c i t e d i n Carbyn 1974). Snow d e p t h i n c r e a s e s f r o m l a t e O c t o b e r t o a maximum o f 40-50 cm i n J a n u a r y o r F e b r u a r y ( T a b l e I ) . T h i s i s f o l l o w e d by r a p i d snowmelt and r u n o f f i n l a t e March and e a r l y A p r i l so t h a t much o f t h e v a l l e y f l o o r i s snow-free by t h e m i d d l e o f A p r i l . Snow a c c u m u l a t i o n v a r i e s c o n s i d e r a b l y a l o n g the v a l l e y (Carbyn 1974) s u c h t h a t t h e a r e a between Henry House and J a s p e r Lake ( F i g . 1) was f r e q u e n t l y s n o w - f r e e i n J a n u a r y o r F e b r u a r y o f 1974-75 and 1975-76. By c o n t r a s t , c o n s i d e r a b l e snow d e p o s i t s were c h a r a c t e r i s t i c o f t h e s o u t h e r n p o r t i o n o f t h e s t u d y a r e a d u r i n g t h e s e months ( T a b l e I ) . V e g e t a t i o n The dominant f e a t u r e o f t h e v e g e t a t i o n i n t h e montane zone (975-1220 m), as d e l i m i t e d by t h e d i s t r i b u t i o n o f d o u g l a s f i r ( P s e u d o t s u g a m e n z i e s i i v a r . g l a u c a ) ( S t r i n g e r and L a R o i 1 9 7 0 ) , i s a c a r p e t o f c o n i f e r o u s e v e r g r e e n f o r e s t c h a r a c t e r i z e d by e x t e n s i v e s t a n d s o f l o d g e p o l e p i n e ( P i n u s c o n t o r t a ) , w h i t e s p r u c e ( P i c e a g l a u c a ) and b l a c k s p r u c e ( P i c e a m a r i a n a ) . I n t e r s p e r s e d Table I. Summary of c l i m a t i c information recorded at Jasper townsite for the period 1973-1976. Temperature (°C) P r e c i p i t a t i o n (cm) Snow accumulation (cm) Mean d a i l y Mean d a i l y Total (rain Month maximum minimum R a i n f a l l Snow equivalent) Jan. -6.8 -16.7 Tr 53.6 4.0 46.9 Feb. -3.2 -13.4 . Tr 15.2 1.3 38.1 Mar. 2.5 -8.5 Tr 19.2 1.6 19.7 Apr. 9.9 -3.2 1.0 5.5 1.4 0.0 May 14.7 " 1.4 2.3 3.6 2.7 0.0 Jun. 18.7 5.4 4.6 0.0 4.6 0.0 J u l . 22.8 7.7 3.5 0.0 3.5 0.0 Aug. 20.6 6.9 3.1 0.0 3.1 0.0 Sep. 18.3 3.1 2.4 0.0 2.4 0.0 Oct. 9.4 -0.4 ' 2.7 9.8 3.4 1.6 Nov. -2.1 -10.9 1.2 43.8 4.5 24.6 Dec. -2.0 -11.2 Tr 34.4 2.4 25.'4 Total 20.8 185.1 34.9 aData obtained from Canadian Department of Transport, Weather O f f i c e . 10 throughout these forests are a number of small grasslands described by Stringer (1969) and i s o l a t e d stands of aspen (Populus tremuloides) and balsam poplar (Populus balsamifera). Various willows (Salix) and alders (Alnus) are common near streams and sedge-grass meadows. Other c h a r a c t e r i s t i c vegetation of t h i s zone are buffaloberry (Shepherdia canadensis), shrubby c i n q u e f o i l ( P o n t e n t i l l a f r u t i c o s a ) , junipers (Juniperus h o r i z o n t a l l s , ^J. communis), s i l v e r b e r r y (Elaeagnus commutata), liearberry (Arctostaphylos rubra) and red os i e r dogwood (Cornus s t o l o n i f e r a ) . Fauna Jasper National Park i s renown for i t s diverse fauna of large mammals which includes seven species of ungulates. Five of these, wapit i , Cervus  canadensis; bighorn sheep, Ovis -canadensis; mule deer, Odocoileus hemionus; moose, Alces americanus; and mountain goat, Oreanmos americanus, are common to abundant, while mountain caribou (Rangifer a r c t i c u s ) and wh i t e - t a i l e d deer (Odocoileus virginianus) are l o c a l l y common and rare r e s p e c t i v e l y . The present population of wapiti i s the r e s u l t of a release of 89 animals i n the Athabasca Va l l e y i n 1920. Wapiti were abundant i n the park h i s t o r i c a l l y but suffered s u b s t a n t i a l mortality as the r e s u l t of severe winters and over harvesting by man. By 1894 t h i s species was absent or at least extremely rare (Soper 1970). W i l d l i f e counts of the wap i t i population by warden service s t a f f i n d i c a t e that f o r the years 1965-1971 the population was stable at about 2,000 animals (minimum number a l i v e ) . W i l d l i f e counts f o r 1972 and 1973 suggested a decline i n numbers to 1400 animals, and the 1974 and 1975 counts indicated a considerable decline to 850 wap i t i f or the park. 11 W i l d l i f e counts i n November 1974-75 and 1975-76 indicated that 302 and 429 wapiti (minimum number a l i v e ) wintered on the main study area. The winter d i s t r i b u t i o n of elk on the study area i s treated i n d e t a i l by Stelfox (1974). In summer, most elk leave the v a l l e y f l o o r with the exception of a herd of females and young i n the area of Rocky River and another which r e g u l a r l y appears i n the Maligne River area. Rocky Mountain bighorn sheep winter i n four main areas of the Athabasca Valley (Fig. 2). W i l d l i f e counts i n November of 1974-75 and 1975-76 suggest that approximately 180-200 sheep winter i n the Athabasca Valley. Mule deer are the most d i f f i c u l t to census. W i l d l i f e counts by the warden service during the period 1966 to 1975 suggest a mean minimum park population of 252 ± 17.5 S . E . animals and v a l l e y population of 166 ± 12.3. There i s some evidence of a decline i n mule deer numbers during the period 1974-1976 s i m i l a r to that observed i n elk and sheep due to the severe 1973-74 winter. Caribou are rare v i s i t o r s to the v a l l e y as are goats. Moose are d i s t r i b u t e d more or le s s evenly along the v a l l e y as habitat permits, however, estimates of t h e i r numbers were not obtained. In ad d i t i o n to coyotes, probably the most p l e n t i f u l large predator i n the park, black (Ursus americanus) and g r i z z l y bears (Ursus arctos) are common residents of the study area i n summer. Wolves are regular v i s i t o r s to the v a l l e y i n winter and are occasionally present i n summer. Red fox, lynx (Lynx lynx), and mountain l i o n ( F e l i s concolor) are present i n low numbers throughout the year. History of the Coyote Population Coyotes have probably inhabited the Athabasca Valley f or hundreds of 12 F i g . 2. Approximate winter d i s t r i b u t i o n of Ovis canadensis i n the Athabasca River Valley, Jasper National Park. 13 14 years. In one of the e a r l i e s t reports of the area (1895), Loring found coyotes abundant i n the f o o t h i l l s outside the present park boundary and shot one specimen near Jasper House (c i t e d i n Soper 1970). Records obtained from Park Superintendent's Annual Reports suggest that coyotes have been abundant to common since the early 1930's. Establishment of the park i n 1907 insured the protection and s u r v i v a l of ungulate populations which had been reduced to low l e v e l s by the combined e f f e c t s of overharvest and severe winters. However, carnivore populations were heavily hunted as a r e s u l t of con t r o l p o l i c i e s thought necessary to insure adequate recovery of ungulates. In 1934 park wardens removed 114 coyotes from the park, many of these from the Athabasca Va l l e y (Fig. 3). Throughout the period 1935 to 1946 an average of 56.1 ± 7.6 coyotes were shot or poisoned annually i n the park. From t h i s period through 1959 coyote numbers were low or cont r o l e f f o r t decreased markedly; 11.5 ± 1.4 coyotes were k i l l e d by wardens annually. Control a c t i v i t i e s appear to have ceased by the l a t e 1950's. The major source of man-related mortality from 1960 to the present i s the r e s u l t of a steady increase i n vehicular t r a f f i c i n the Athabasca Va l l e y (Table I I ) . From 1970 to 1976, an average of 7.7 ± 1 . 2 coyotes were k i l l e d annually on the study area by ve h i c l e s . Thus, the population has not been hunted since the l a t e 1950's, however, a l e v e l of man-related mortality p e r s i s t s . 15 F i g . 3. Number of coyotes k i l l e d i n Jasper National Park during periods of predator c o n t r o l ( • ) and by motor vehicles ( o ) from 1934 to 1976. 16 100 2 50 o V) cn o •o 0) *-* o >» o o a> x> E 1935 1945 1955 T i m e 1965 1975 / 17 Table I I . Summary of v e h i c u l a r t r a f f i c i n a Jasper N a t i o n a l Park, 1960 to 1971. Number of v e h i c l e s e n t e r i n g 1960-61 91,357 1961-62 95,239 1962-63 139,813 1963-64 137,499 1964-65 149,960 1965-66 161,412 1966-67 194,989 1967-68 212,946 1968-69 284,966 1969-70 401,660-1970-71 470,106 S t a t i s t i c s s u p p l i e d by V i s i t o r S e r v i c e s , Jasper N a t i o n a l Park. \ 18 3 GENERAL METHODS In t h i s section, I describe the general methodology used to study the coyote population. S p e c i f i c methods are presented i n subsequent chapters. Coyotes were trapped using Oneida-Victor No. 2 C o i l Spring traps. To minimize the in j u r y to captured coyotes, I t i g h t l y wrapped the trap jaws with several layers of canvas. Care was taken to deodorize traps i n a sol u t i o n of logwood c r y s t a l s and spruce sap before use. This treatment markedly increased capture success by elminating human odour. Of the several d i f f e r e n t trap sets used, the food-cache set described by Israelson (1968) was most successful. B r i e f l y , the method involves digging a 20-25 cm deep c o n i c a l hole into which a small amount of commercial b a i t (Northwoods, Thief River F a l l s , Minnesota) i s placed. The trap i s concealed i n the fan of earth t a i l i n g s produced by the hole construction. The trap i s anchored to a drag log or tree or may be equipped with a drag hook. I found that a strong spring placed between the trap and the drag log appeared to reduce or eliminate i n j u r y . During each trapping period, I maintained 12 to 24 food-cache sets. Each set was checked d a i l y , usually i n the morning, but sets were often checked twice d a i l y . When a coyote was captured the set was immediately re-established. I restrained trapped coyotes by placing an adjustable loop of rope, attached to one end of a long pole, over the animal's head. I then muzzled, bound the legs, and released the animal from the trap. Coyotes captured i n the f a l l were e a s i l y handled without the use of drugs. I assumed t h i s was due to lower l e v e l s of c i r c u l a t i n g testosterone outside the breeding season. In the spring, coyotes were f a r more aggressive and were given an i n t r a -19 muscular i n j e c t i o n of equal parts of phencyclidine hydrochloride (Sefnylan, 100 mg per cc, Bio-Ceutics Laboratories, Inc.) and acepromazine maleate (Atravet, 25 mg per cc, Ayerst Laboratories) as described by Seal and Erickson (1969). Dens suspected of housing coyote pups were excavated i n May or early June. Pups were measured, tagged, and returned to an unexcavated section of the den as soon as p o s s i b l e . To insure pups would not stray before the return of t h e i r parents, s t i c k s or pieces of sod were placed at the tunnel entrance. In no instance were pups abandoned a f t e r our disturbance. Coyotes greater than 2 kg were weighed on a spring scale to the nearest 0.2 kg. Pups were weighed to the nearest 25 g on a 2 kg Pesola. Body length, t a i l length (from f i r s t to l a s t caudal vertebrae), ear length, hind foot length and neck circumference were recorded f o r each animal. I also examined the animal f o r ecto-parasites, wounds or other i n j u r i e s , and noted the general condition of the pelage. Reproductive condition of males was assessed by noting the s i z e and p o s i t i o n of the testes. In females, the colour of the vulva, colour and s i z e of t i t s , and presence of palpable fetuses were used to assess reproductive status. I determined the age of coyotes from the patterns of wear on the i n c i s o r s and canine teeth (Gier 1968, unpubl. data). To check the r e l i a b i l -i t y of t h i s method, 30 coyote carcasses were aged f i r s t by the tooth wear method and then by counting cementum annuli of canines and f i r s t lower premolars (Linhart and Knowlton 1967). The r e - c a l i b r a t e d scale (Appendix I) was then applied to correct the age of captured adults. Also presented i n Appendix I i s a re-evaluation of age determination i n coyotes by the cementum annuli method. Juvenile coyotes were aged using the growth 20 regressions of Gier (1968), tooth eruption patterns and age of eye opening (Bekoff and Jamieson 1975). A l l adults and subadults (6-12 months of age) were equipped with a radio transmitter i n the form of a c o l l a r (Fig. 4). The radio c o l l a r weighed approximately 320 g or 2.5 percent of the coyote's adult body weight. Radios transmitted a pulsed s i g n a l on one of 12 frequencies near 30 MHz. By using d i f f e r e n t periods of the pulsed s i g n a l more than one animal could be distinguished on a si n g l e frequency. I used period of 60, 80, and 120 beats per min. D e t a i l s of the e l e c t r o n i c components and a c i r c u i t diagram of the transmitters are given i n Seidensticker et a l . (1970). The transmitted s i g n a l was picked up by a tuned, magnetic, dipole loop or a whip antenna; The receiving range of the two antennae on the ground was 2.4 and 4.0 km f o r the loop and whip r e s p e c t i v e l y . I used the whip antenna to i n i t i a l l y locate an animal and the dipole loop to obtain d i r e c t i o n a l compass bearings. The l o c a t i o n of a r a d i o - c o l l a r e d coyote, was usually determined by t r i a n g u l a t i o n . D i r e c t i o n a l compass bearings were plotted on 1:50,000 topographic maps of the study area and the i n t e r s e c t i o n 2 of two or more bearings referenced on a .25 km acetate g r i d . In general, I attempted to obtain one radio l o c a t i o n per animal per day. In ad d i t i o n to radios, a l l subadults and adults were ear tagged and given a numbered, coloured, p l a s t i c neck tag (Fig. 4). Juveniles were eartagged only. Ear tags consisted of a numbered, colour-coded, Rototag (Nasco) to which was added a 5 cm coloured, p o l y v i n y l streamer. Neck tags (Ketchum Manufacturing) enabled rapid i d e n t i f i c a t i o n of coyotes at a distance of 1 km or more using a 15 to 40 power spotting scope. Detailed drawings of the pelage of a l l coyotes i n the population which were r e g u l a r l y observed allowed the i d e n t i f i c a t i o n of many i n d i v i d u a l s which 21 F i g . 4 . A l l subadults and adult coyotes captured were equipped with a radio transmitter i n the form of a c o l l a r and were given an ear tag and neck tag. i 23 I was unable to capture. Two examples of these composites are given i n F i g . 5. V a r i a t i o n i n the colour and d i s t i n c t n e s s of the shoulder saddle, precaudal spot, and hip s t r i p s were the most useful pelage c h a r a c t e r i s t i c s . I divided the year into a f i v e month summer (May 1 to September 30) and a seven month winter (October 1 to A p r i l 30). This d i v i s i o n corresponds with the phenology of the Columbia ground s q u i r r e l (Spermophilus columbianus) a major summer prey of the coyote and with the presence and absence of snow on the study area. The standard error (S.E.) i s given with means, unless stated otherwise. 24 F i g . 5. Drawings of coyote pelage c h a r a c t e r i s t i c s used to i d e n t i f y c e r t a i n members of the population. 26 4 CAPTURE DATA AND TELEMETRY STUDIES Capture Data Forty-three coyotes c o n s i s t i n g of 10 adult males, 10 adult females, s i x ju v e n i l e males and 16 j u v e n i l e females were captured 51 times (Table I I I ) . Two adult females were recaptured. C4, f i r s t captured on November 3, 1974, was recaptured approximately s i x months l a t e r on A p r i l 13, 1975, 1.9 km from the i n i t i a l capture s i t e . C15 was recaptured four months l a t e r on May 13, 1975, 2.6 km from her f i r s t capture l o c a t i o n . Six ju v e n i l e s were recaptured, four at dens (caught by hand) and two i n traps. Both animals recaptured i n traps, one male and one female, were captured 38 days l a t e r less than 2.0 km from t h e i r i n i t i a l capture s i t e . In addition to 43 captured animals, weight and body measurements were taken from 29 r o a d - k i l l e d coyotes. These data for adults are presented i n Appendix I I . Mean body measurements and mean weight of 20 adult females were: body length, 88.4 ± 1.42 cm; t a i l length, 32.5 ± 0 . 5 3 cm; hind foot, 18.8 ± 0.13 cm, and 11.5 ± 0.33 kg. For 19 adult males, mean body measurements and mean weight were as follows: body length, 89.4 ± 1.81 cm; t a i l length, 33.7 ± 0.51 cm; r i g h t hind foot, 18.8 ± 0.22 cm and 12.1 ± 0.45 kg. The heaviest female and male weighed 13.6 and 15.0 kg r e s p e c t i v e l y . Telemetry Studies Radio telemetry provided information on a number of aspects of coyote biology including 1) reproduction, 2) the s i z e , u t i l i z a t i o n , and d i s t r i b u t i o n of home ranges, 3) movements incl u d i n g d i s p e r s a l , 4) a c t i v i t y , and 5) the nature and frequency of coyote associations. As these topics are dealt with Table I I I . Results of the capture program i n Jasper National Park, 1974-1976. Number of coyotes radio tagged i s given i n parentheses." Number Number Total number captured recaptured captured Capture period Adults Young Adults Young F a l l , 1974 6(6) . 3(3) 0 0 9 Spring, 1975 9(9) 13 l a 3 26 ' Summer, 1975 0 4 0 1 5 F a l l , 1975 0 3(3) l a 2 6 Spring, 1976 5(5) 0 0 0 5 Total no. ' 20(20) 23(6) 2 6 51 Animal equipped with a new radio transmitter. 28 i n d i f f e r e n t chapters of the thes i s , t h i s section serves as a b r i e f summary of the information c o l l e c t e d . During the period September, 1974 to February, 1977, I obtained 4,736 radio locations of 26 d i f f e r e n t coyotes (Table IV). The radios of f i v e coyotes, three adult females and two adult males, transmitted f o r an average of 484.6 days, ranging from 427 to 618 days. Six radio c o l l a r s transmitted for an average of 243.5 ± 15.88 days, a l l functioning greater than s i x months but less than one year; F i n a l l y , 15 radios operated f o r les s than s i x months (x = 69.5 ± 11.05 days). Of t h i s l a s t group, three radios malfunctioned, three r a d i o - c o l l a r e d coyotes dispersed, four animals died and f i v e radios were operating normally when study was terminated s h o r t l y a f t e r coyotes were equipped with transmitters. Radio-tagged coyotes were not always located. The a b i l i t y to locate an animal was a function p r i m a r i l y of the s o c i a l class to which the animal belonged. Resident adult members of a p a i r or pack were con s i s t e n t l y easier to locate; 94.0 percent of attempts resulted i n a successful radio l o c a t i o n . However, only 73.4 percent of attempts resulted i n radio locations of dispersing coyotes or s o l i t a r y residents. Combining these groups, an average of 86.7 ± 6.32 percent of attempts resulted i n successful radio l o c a t i o n s . 29 Table IV. Summary of radio telemetry data for 26 coyotes captured i n Jasper National Park. Date Date of No. of days No. of Animal radio tagged l a s t s i g n a l transmitting radio lc C1,Y0Y M a 24 09 74 b 12 11 74 47 92 C2, Ad F 09 10 74 20 01 75 100 104 C3, Ad F 29 10 74 01 07 75 244 320 C4, Ad F 03 11 74 22 07 76 c ,618 697 C5, Ad F 05 11 74 16 12 74 41 36 C6, Ad F 11 11 74 25 06 75 225 305 C7, YOY M 13 11 74 04 02 75 84 63 C8, YOY M 13 11 74 22 11 74 9 9 CIO , Ad M 13 11 74 06 03 76 470 513 C l l Ad F 06 04 75 22 07 76 472 508 C12 , Ad M 16 04 75 19 08 75 125 261 C13 , Ad M 26 04 75 27 10 75 184 320 C14 , Ad M 26 04 75 29 05 75 33 26 C15 , Ad F 13 05 75 22 07 76 436 290 C16 i Ad M 29 04 75 10 06 75 42 35 C17 , YOY F 19 09 75 12 05 76 236 120 C18 , YOY M 20 09 75 29 06 76 283 181 C19 , Ad M 03 05 75 10 06 75 36 48 C20 , Ad M 12 05 75 22 07 76 427 429 C22 , YOY F 29 10 75 27 03 76 149 73 C23 , Ad M 10 04 76 22 07 76 102 65 C24 , YOY F 14 04 76 22 07 76 99 60 C25 , Ad M 17 04 76 22 07 76 95 65 C26 , YOY F 28 04 76 11 02 77 289 56 C27 , Ad F 03 05 76 22 07 76 80 53 C50 , Ad M 12 04 75 16 04 75 4 7 YOY = young of year, M = male Day, month, year Date study was terminated ) 30 5 POPULATION BIOLOGY Coyotes occur at r e l a t i v e l y low d e n s i t i e s . They are crepuscular or l a r g e l y nocturnal i n many areas and are highly mobile. These c h a r a c t e r i s t i c s present s p e c i a l sampling problems which make an accurate determination of numbers d i f f i c u l t . Consequently, indices of abundance are commonly used. ( rather than an.estimate of absolute numbers (see Linhart and Knowlton 1975 for review). Despite these d i f f i c u l t i e s I attempted a t o t a l enumeration of the population. I f e l t the s i z e and shape of the study area would lend i t s e l f n i c e l y to th i s approach. Summer estimates of density are based on observa-tions of known i n d i v i d u a l s and t h e i r associates - and the number and s i z e of known l i t t e r s . Density estimates i n winter are based on t o t a l counts of known i n d i v i d u a l s and t h e i r associates attending ungulate carcasses p o s i t i o n -ed throughout the study area. These estimates represent the minimum number a l i v e during each season and are thought to r e l i a b l y estimate t o t a l numbers. Numbers I estimated that approximately 55 coyotes were on the study area during July of 1975 and 1976 (Fig. 6). In July, 1974, I had j u s t a r r i v e d on the study area and was unable to obtain complete information on the number of l i t t e r s or the t o t a l number of adults. About 40 coyotes were on the study area i n March, 1975 and 1976. These data suggest a post-whelping (July)' 2 density of 0.46 coyote per km and a l a t e winter (March) density of 0.35 2 coyotes per km . Gier (1968) reported a post-whelping densityvof 0.8 2 coyotes per km f o r Kansas. S i m i l a r l y , Knowlton (1972) estimated a summer 2 2 density of 0.9 coyotes per km on a 48 km area i n south Texas. Working i n 31 F i g . 6. Minimum number of coyotes a l i v e i n J u l y and March, 1974-1976, i n the Athabasca R i v e r V a l l e y . v Data a v a i l a b l e f o r only f i v e of nine coyote groups. ^The l i t t e r of three groups was not found, however, I assumed two pups a l i v e per group at the time of the census. 32 1974 1975 1976 S e a s o n 33 an aspen-fir forest i n north-central Minnesota, Chesness and Bremicker (1974, paper presented at Coyote Research Workshop, Denver, Colorado) reported 2 post-whelping de n s i t i e s of 0.2 to 0.4 coyotes per km more s i m i l a r to those found i n Jasper. N e l l i s and Keith (1976) estimated winter de n s i t i e s of 0.1 2 to 0.6 coyotes per km for a mixed farming-boreal forest region i n c e n t r a l Alberta. The highest f a l l d e n s i t i e s are those reported by Knowlton (1972) 2 of 1.5 to 2.3 coyotes per km i n south Texas. My winter estimate f a l l s 2 within the range of coyote de n s i t i e s (0.2 to 0.4 coyotes per km ) considered by Knowlton (1972) to be t y p i c a l over much of i t s range. Based on these data i t appears that coyotes i n Jasper occur at average d e n s i t i e s . To determine the r e l i a b i l i t y of my winter density estimate, I p l o t t e d the percentage of known coyotes observed against time. If I knew the majority of animals i n the population t h i s p l o t would asymptote near 100.0 percent. In both the winters of 1974-75 and 1975-76 the percentage of known coyotes observed asymptoted i n January at about 90.0 percent (Fig. 7). The r e l a t i o n s h i p asymptoted at le s s than 100.0 percent because some observations of coyotes were at distances too great or 'for periods too short to i d e n t i f y the animal. Therefore, some observations of known coyotes were probably scored as unknown. I t i s also l i k e l y that there were several coyotes i n the populations of which I was unaware. I conclude that my estimate of absolute population s i z e i s reasonably accurate. Reproduction Information on coyote reproduction was gathered i n the spring and summer of 1974 through 1976. In 1974, s i x l i t t e r s were born on the study area (Table V), but t h i s was an incomplete record f o r reasons described 34 F i g . 7. Percentage of t o t a l s i g h t i n g s which were of known coyotes as a f u n c t i o n of months during the w i n t e r of 1974-75 ( O ) and 1975-76 ( • ). Number as s o c i a t e d w i t h each symbol . represents the t o t a l number of s i g h t i n g s per month. 35 Oct Nov Dec J a n Feb Mar 36 Table V. Summary of coyote l i t t e r s born on the study area from 1974 to 1976. Minimum number born Name of l i t t e r 1974 1975 1976 Rocky R i v e r 4 ( C 6 ) a none(C6) 5(C6) Talbot Lake UNK b(C3) 4(C3) ; none C i n q u e f o i l UNK(?) 2(DIS) UNK(DIS) Snaring K ? ) 2(ALI) 3(ALI) C o l i n 2(BLB) 2(BLB) NA C(BLB) Pa l i s a d e s UNK(?) 6(C15) NA(SAL) Athabasca 4(C11) l ( C l l ) 5(C11) Maligne 4(C2) 6(TES) 4(TES) M i e t t e 2(?) 5(?) UNK(?) T o t a l no. young 17 28 17 aCode i n brackets i n d i c a t e s female parent. A question mark i n d i c a t e s female's i d e n t i t y u n c e r t a i n . UNK - no evidence of l i t t e r found. NA - number of young unknown. 37 e a r l i e r . L i t t e r sizes given i n Table V represent the minimum number of young that I knew were born. They are biased data as only a small number of l i t t e r s were enumerated at the den sh o r t l y a f t e r b i r t h . Therefore, these data should not be used as an estimate of l i t t e r s i z e . Seventeen young were a l i v e on the area i n J u l y , 1974. I was not able to f i n d the n a t a l or rearing dens i n the Talbot, Cinq u e f o i l and Palisades ranges though they are believed to have been present (Fig. 13). One pup was captured i n the Snaring range i n September before d i s p e r s a l generally takes place and a recently used den was located near the capture s i t e . In 1975, eight l i t t e r s t o t a l l i n g 28 young were born on the study area (Table V).. Of the paired, adult females i d e n t i f i e d , only female C6 did not produce a l i t t e r i n 1975. She was c l o s e l y monitored by radio during the period of January-May and frequently observed, however, at noi^ time was there evidence of a pregnancy. C6 produced at l e a s t 4 young i n 1974 and a l i t t e r of f i v e i n 1976. As she had the same mate i n both 1975 and 1976 and l i k e l y i n 1974, she must have aborted the pregnancy at an early stage. The l i t t e r i n the Snaring range was not located during the summer, however, observation of t h i s pack i n the winter of 1975-76 suggests that at l e a s t two pups were born. In 1976, the data were again incomplete. Six l i t t e r s were born c o n s i s t i n g of 17 young (Table V). The Talbot Lake range no longer existed as female C3 was k i l l e d i n 1975 and her mate dispersed. I was unable to f i n d evidence of a l i t t e r i n the Miette or Cinquefoil ranges, however, I l e f t the study area i n l a t e July before the young are highly mobile. I t i s l i k e l y a l i t t e r was produced i n both areas. A f r e s h l y used den was located i n both the Palisades and C o l i n ranges but the number of young was unknown. 38 Female C15 f a i l e d to produce a l i t t e r i n 1976. She became a s o l i t a r y r e s i -dent i n the winter of 1975-76 and although she established a new home range in the Maligne Valley ( F ig. 17) she did not obtain a mate. She was paired i n February, 1977, but I do not know i f she produced a l i t t e r that spring. The best estimate of coyote l i t t e r s i z e for t h i s l a t i t u d e i s 5.3 (n=26) given by N e l l i s and Keith (1976) for c e n t r a l Alberta. The reproduc-t i v e t r a c t s of four females were c o l l e c t e d between March and July i n t h i s study. Seven embryos were present i n one 5 year old female and 4, 5, and 6 placental scars were c l e a r l y v i s i b l e i n the other three. N e l l i s and Keith (1976) found an average of 6.0 scars per t r a c t . Knowlton (1972) found that averages ranged from 5.1 to 6.9 scars per t r a c t for d i f f e r e n t areas of Texas and Hamlett (1938) reported 5.1 and 6.7 scars per t r a c t f o r western United States. The date of p a r t u r i t i o n was estimated f o r eight l i t t e r s ; s i x were determined by regression of body length on age (Gier 1968) to age the pups and two were estimated from marked changes i n the a c t i v i t y of radio-tagged females. P a r t u r i t i o n occurred from March 29 to June 10. Five of eight l i t t e r s were born from March 29 to A p r i l 12, two from May 1 to May 3 and one i n June. The dates of p a r t u r i t i o n of female coyotes i n Jasper are within the range of those examined by Hamlett (1938) , Gier (1968) and Gipson (1972). M o r t a l i t y T r a f f i c deaths accounted for 72.7 percent of known summer coyote m o r t a l i t i e s and 77.3 percent of known winter deaths (Table VI). In summer, Table VI\ Number of coyote m o r t a l i t i e s i n the Athabasca River Valley during the period 1974-1977. Summer 1974 Winter 1974-75 Summer 1975 Winter 1975-76 Summer 1976 Winter 1976-77 Source YOY Adults YOY Adults YOY Adults YOY Adults YOY Adults YOY Adults Vehicles 3 1 1 6 2 1 3 2 1 5 Predation 1 1 Disea'se 1 l a Unknown 1 2 1 Total 4 11 5 4 2 7 aSuspected as a contributing f a c t o r . 40 the mean number of coyote deaths from a l l known sources was 4.7. In winter, an average of 7.7 coyotes died annually from 1974-75 to 1976-77. The high proportion of man-related deaths i s p a r t i c u l a r l y important as the study population was selected on the assumption that i t was minimally influenced by man. Predation or i n t e r s p e c i f i c aggression accounted for two coyotes deaths. An adult female, C3, was k i l l e d by a mountain l i o n i n July, 1975, several hundred meters from an elk carcass. She had a f u l l stomach of f r e s h l y consumed elk meat at the time of her death. Although most of the animal had been consumed, the head, v i s c e r a , and the one remaining hind leg were buried i n a manner t y p i c a l of mountain l i o n s . I t i s l i k e l y that t h e . l i o n surprised the coyote feeding on the carcass or leaving the area a f t e r feeding. The second incidence, reported to me by park wardens, occurred on January 1, 1977. A coyote foraging within the fence of the sanitary l a n d f i l l was attacked and k i l l e d by a wolf. A necropsy indicated that a si n g l e canine tooth puncture of the l e f t p a r i e t a l bone was the cause of death. Three i n c i s o r impressions were also found i n the r i g h t p a r i e t a l near the s a g i t t a l c r e s t . Substantial hemorrhaging occurred i n the head and neck area before death. That none of the coyote was consumed suggests i n t e r s p e c i f i c aggression rather than an act of predation. F. Burstrom ( c i t e d i n Cowan 1947) reported an incident where four wolves had caught and k i l l e d a coyote i n the Athabasca V a l l e y near Devona. Carbyn (pers. comm.) has reported several s i m i l a r aggressive attacks by wolves on coyotes. Although there are recurrent reports of t h i s nature, I believe wolf-induced m o r t a l i t y to be rare. 41 Sarcoptic mange (Sarcoptes scabiei) was responsible f o r the death of one adult male and possibly was a contributing factor i n the death of another. RK17 was discovered i n a coyote den i n May,. 1975, i t s body l a r g e l y devoid of hai r i n a pattern c h a r a c t e r i s t i c of mange (Samuels pers. comm.) (Fig. 8). L i t t l e h a i r was found i n the den i n d i c a t i n g that the animal had sought shelter from winter temperatures a f t e r l o s i n g much of i t s i n s u l a t i n g coat. Male C16 showed evidence of mange throughout the winter of 1974-75. Observations from a distance indicated h a i r loss on both hips and the area about the base of the t a i l . Pinkish crusted les i o n s i n these areas provided further evidence of skin i n f e c t i o n . When the animal was captured on A p r i l 29, 1975, I found a considerable number of crusted les i o n s and scar t i s s u e on the hips and rump area. On June 10, 1975, C16 was found dead about 5 km south of i t s former range; i t had apparently begun to disperse. There was evidence that the mange i n f e c t i o n had worsened. The t a i l was badly scarred and devoid of h a i r and there were crusted, coalesced lesi o n s on the hips and hind legs. Unfortunately, the animal had probably been dead f o r several days. When found much of the trunk had been consumed, the cranium showed four canine punctures approximately coyote s i z e and the r i g h t side of the cranium was missing. Thus, mange may not have been the proximate cause of death but was l i k e l y a contributing f a c t o r . During the study mange was not common. Only f i v e of 43 (11.6 percent) coyotes captured showed evidence of sarcoptic mange. In none of these animals did I attempt to i s o l a t e mites. Thus, I can only i n f e r that mange was involved. However, the l o c a t i o n of crusted les i o n s and hai r l o s s i n a l l cases agreed with the pattern described for coyotes infected with S_. ( 42 F i g . 8. Pattern of h a i r loss which i s c h a r a c t e r i s t i c of advanced scarcoptic mange i n f e c t i o n . 4 3 44 s c a b i e i (Samuels pers. comm.)- An adult female observed only once i n October, 1974, was severely infected with mange. Crusted le s i o n s were located over much of the animal's body. The midline of the back and neck, and the .rostrum were the only areas escaping s u b s t a n t i a l h a i r l o s s . Three members of the Athabasca Pack captured i n the spring of 1975 apparently had mange. In each case the i n f e c t i o n appeared to wane as the summer progressed and a f t e r the summer molt a l l three seemed to have f u l l y recovered from the disease. Mange symptoms did not reappear during the winter of 1975-76. Drowning claimed one adult female i n January, 1975. Radio bearings indicated that she was lodged beneath the i c e of the Athabasca River at the edge of her t e r r i t o r y . I observed t h i s female several days before her death at which time I detected nothing abnormal i n her behaviour. I t i s l i k e l y that she broke through the i c e while crossing or t r a v e l l i n g along the r i v e r . F i n a l l y , three coyotes, one adult female and two young died of unknown causes. I used the method of Trent and Rongstad (1974) to c a l c u l a t e coyote s u r v i v a l and m o r t a l i t y rates from radio-telemetry data. Su r v i v a l over n days i s given by: where x i s the number of radio-coyote days and y i s the number of deaths observed over n days. A radio-coyote day i s 1 radio-tagged coyote i n the f i e l d for 1 day. Annual s u r v i v a l rate was calculated as the product of quarterly s u r v i v a l rates. M o r t a l i t y rate i s 1-Sn. Nine of 26 radio-tagged 45 coyotes died between September, 1974 and July, 1976. Annual m o r t a l i t y of subadult and adult coyotes over t h i s period was 0.45 ± 0.225 (n=2). Note, however, there i s considerable v a r i a t i o n between years. Knowlton (1972) stated that annual m o r t a l i t y rates of yearlings and older coyotes may exceed 40 percent even i n unexploited populations. The estimated annual m o r t a l i t y i n Jasper, however, i s s i m i l a r to that reported i n a heavily hunted population by N e l l i s and Keith (1976). They estimated an annual mortality of juveniles and adults of 0.40. In other hunted populations, annual mortality of yearlings and adults may average 0.62 ± 0.200 (n=2) (Knudsen 1976). Given the v a r i a t i o n i n annual m o r t a l i t y between years i n Jasper and other areas (see Knudsen 1976), i t i s only possible to state that the estimates i n Jasper are among the lowest reported. 46 6 HOME RANGE AND PATTERNS OF SPACE USE Animals frequently l i m i t t h e i r a c t i v i t y to a p a r t i c u l a r area within which they f i n d the resources necessary f o r growth, maintenance, and repro-duction. Such an area i s defined as a home range (Burt 1943). Size and shape are the home range parameters most frequently estimated (see S t i c k e l 1954, Sanderson 1966, Van Winkle 1975, f o r reviews). Recently, however, the u t i l i z a t i o n d i s t r i b u t i o n (Hayne 1949) which describes the d i s t r i b u t i o n of an animal's p r o b a b i l i t y of occurrence at various points i n space, has received serious attention (Ford and Krumme i n press, Van Winkle 1975). Often these home range parameters have been investigated f o r t h e i r putative i n t r i n s i c value as i f by themselves these properties increase our understand-ing of the use of space by animals. Whereas, home range s i z e and shape may be of l i m i t e d value to the manager, they are of l i t t l e general i n t e r e s t i n themselves. However, when coupled with e c o l o g i c a l and behavioural informa-t i o n , the s i z e , shape and u t i l i z a t i o n d i s t r i b u t i o n of an animal's home range become meaningful b i o l o g i c a l parameters. In the present study, coyote home ranges are of i n t e r e s t to the extent that they reveal (1) the d i s t r i b u t i o n of i n d i v i d u a l s i n time and space, (2) the r e l a t i o n s h i p between home range s i z e and coyote-group s i z e , and (3) u t i l i z a t i o n of space by coyotes i n response to s o c i a l and e c o l o g i c a l f a c t o r s . Methods I used the minimumj.convex polygon method (Mohr 1947) to delimit the boundaries of coyote home ranges. I modified the method to include only 47 the 95 percent of the radio relocations c l o s e s t to the range center. This recognizes that s a l l i e s from the area most frequently used should not be included i n the home range (Burt:1943, H i b l e r 1976). To determine whether an animal had a home range, I plo t t e d an index of area u t i l i z e d against the number of radio locations per coyote. This index was derived by counting the cumulative number of grids (see General Methods) enclosed by a convex polygon j o i n i n g the outermost locat i o n s as each new l o c a t i o n was added. Home range was achieved when an increase i n the index was _£2.5 percent for a period of one month or 30 radio l o c a t i o n s , whichever was l e s s . Only asymptotic r e l a t i o n s h i p s were denoted as a home range. Home range siz e was determined using a compensating polar planimeter. Each range was measured twice and the mean value reported. Results Home range as the asymptote of the observation-area curve As the number of observations of an animal accumulates the area where i t has been found increases asymptotically (Hayne 1949, S t i c k e l 1954, Sanderson 1966). The r e l a t i o n s h i p i s termed the observation-area curve (Odum and Kuenzler 1955) and i t s asymptote, as stated above, i s an estimate of home range s i z e . However, another curve i s possible and i s l i k e l y to occur for subadults. In t h i s case the area i n which an animal has been observed increases s t e a d i l y with time. As they represent very d i f f e r e n t s trategies of space use, i t i s important to d i s t i n g u i s h between these two curves. Unfortunately, few authors take the time to determine the shape of t h i s r e l a t i o n s h i p f o r i n d i v i d u a l animals. This makes i t d i f f i c u l t to sort 48 out the r e a l influence of b i o l o g i c a l v a r i a b l e s on home range s i z e from measurement error associated with i n s u f f i c i e n t data or the measurement of a d i f f e r e n t phenomenon (e.g. d i s p e r s a l ) . In t h i s study, 17 of 26 radio-tagged coyotes had a home range. The index of area u t i l i z e d of nine radio-tagged coyotes did not asymptote based on radio locations alone (Table VII).' C19 and C26 were known to have used the same home range for at le a s t one year based on observations. One resident female died before s u f f i c i e n t data were a v a i l a b l e and s i x coyotes dispersed. The number of radio locations and number of days required to reach an asymptote i s presented i n Table VII. Daily radio locations were attempted, but not achieved for a l l coyotes. Nevertheless, for 10 females and f i v e males, radio locations were obtained approximately d a i l y . For these coyotes, there was no s i g n i f i c a n t difference i n the number of locations required to reach an asymptotic index of area u t i l i z e d between males (41.2 ± 10.83 locations) and females (41.9 ± 5.67 locations) (t = 0.07, d.f. = 13, P >0.05). These radio locations were obtained over a period of 44.1 ± 8.10 days for females and 46.8 ± 14.03 days for males. Combining males and females, an average of 41.7 ± 5.00 locations over a period of 45.0 ± 6.86 days were necessary to define the s i z e of coyote home ranges. Two males, C10 and C18, were not included i n the above comparison. C10, unlike other males i n the sample, was a s o l i t a r y resident whose home 2 range (>65 km ) was considered a t y p i c a l l y large. Thus, 113 radio locations i n 163 days were necessary to determine the s i z e of ClO's home range. The second male (C18) excluded from the sample, underwent a period of d i s p e r s a l before becoming resident. In t h i s case, 98 locations i n 163 days defined 49 Table VII. Number of radio locations and days required to reach an asymptotic index of. area u t i l i z e d for 26 coyotes i n Jasper, Alberta. No. f i x e s to No.days to Duration of Cumulative no. of Animal Age Sex asymptote asymptote asymptote (days) f i x e s at successive asymptotes CI YOY M - - None -C7 YOY F 62 83 140 62 C8 YOY M - . - • None -C17 YOY F - - None -C18 YOY M 98 163 118 98 C22 YOY F 30 34 66 30 C24 YOY . F 22 24 75 22 C26 YOY F - - None c _ C2 Ad F 47 23 ' 77 47 C3 Ad F 14 12 35 14 7 19 63 56 1 1 114 92 C4 Ad F 40 40 135 40 37 27 71 170 1 1 339(534) 237 C5 Ad F - - None d -C6 Ad F 62 83 140(521) 62 C l l Ad F 35 28 54 35 41 45 344(713) 124 C15 Ad F 68 69 74 68 88 167 104(299) 220 C27 Ad F 39 45 35 39 CIO Ad M 113 163 312 113 C12 Ad M 25 24 49 25 1 1 45 55 1 1 30(736) 94 C13 Ad M . 18 19 43 18 1 1 90(233) 52 C14 Ad M • - - None -C16 Ad M - - None -C19 • Ad M - - None e(801) -C20 Ad M 80 98 179 80 C23 Ad M 37 43 59 37 C25 Ad M 46 50 45 46 C50 Ad M - - None -a B a s e d on r a d i o - t e l e m e t r y d a t a a l o n e . The number i n p a r e n t h e s e s g i v e s t o t a l d u r a t i o n o f asymptote based on s i g h t i n g s a f t e r t h e r a d i o s t o p p e d . . E n t r i e s w h i c h a r e u n d e r s c o r e d i n d i c a t e t h a t t h e a n i m a l d i e d . °Study t e r m i n a t e d b e f o r e s u f f i c i e n t number of r a d i o l o c a t i o n s o b t a i n e d , b ut known to have a s y m p t o t i c range f r o m p r e v i o u s s i g h t i n g s d u r i n g w i n t e r 1975-76. ^ D i e d a f t e r 41 days and 36 f i x e s . e R a d i o f a i l e d a f t e r 48 f i x e s i n 36 d a y s , but -known t o have a s y m p t o t i c range f r o m subsequent . . s i g h t i n g s . 50 the home range s i z e . The above analysis applied only to the f i r s t asymptote between area u t i l i z e d and time. More than one asymptote was evident f o r s i x coyotes, four females and two males (Table VII). Female C3 held a home range which increased i n s i z e on three occasions (Fig. 9). Her i n i t i a l home range was u t i l i z e d u n t i l December 16, 1974, a period of 35 days. Over the span of several weeks the area covered by th i s female increased 39.2 percent. The si z e of her home range was stable for the next 63 days. On March 8, 1975, her range increased a f i n a l 18.2 percent and was stable thereafter u n t i l her death i n July, 1975. S i m i l a r l y , three asymptotes are evident i n the rad i o - l o c a t i o n data of female C4 (Fig. 10). This y e a r l i n g female occupied the same area for 135 days from December, 1974 to l a t e A p r i l , 1975. At th i s time she began t r a v e l l i n g with the Athabasca Pack. Concurrently, her range increased 61.2 percent i n 10 days. On June 8, 1975, 26 days l a t e r , her home range increased 6.4 percent. She occupied t h i s larger area f o r 71 days. F i n a l l y , on August 14, 1975 her home range increased by 18.0 percent, an area which contained a l l . subsequent radio locations of t h i s female u n t i l the ' termination of the study i n July, 1976. The area within which I consistently located female C l l was constant from early May, 1975, to l a t e June, 1975, a period of 54 days. From l a t e June through early July her index of area u t i l i z e d increased 48.6 percent. This l i k e l y corresponded with the period of increased mobility of her l i t t e r . Subsequent-l y , C l l ' s index of area u t i l i z e d was constant f o r 344 days. In October, 1975, female C15 dispersed from her home range. Approximately f i v e months l a t e r she established a new home range which was held f or 104 days. Increas-es i n the home range s i z e of males C12 and C13, both members of the '51 Fi g . 9. Relationship between an index of area u t i l i z e d and time for adult, female C3 from October, 1974 to July, 1975. 53 F i g . 10. Relationship between an index of area u t i l i z e d and time for adult, female C4 from October, 1974 to July, 1976. Number of radio locat ions (log scale) 55 Athabasca Pack, coincided with an increase i n the home range of th i s pack's alpha female, C l l . Home range f i d e l i t y Resident adult coyotes occupied the same home range f o r considerable periods of time (Table VII). In Table VII, entries which are underscored indicate that the coyote died; otherwise the values represent the period, from capture u n t i l the end of the study, that an animal u t i l i z e d the same area. Seven radio-tagged coyotes, four females and three males, occupied the same home range f o r periods ranging from 299 to 801 days. The longest period of residence on the same range was 1,094 days (-3 years) by female C l l . She was f i r s t observed on July 26, 1974 and l a s t seen on Ju l y 25, 1977, 1.5 km from the i n i t i a l s i t e of observation. In addition to marked coyotes, eight resident adults, i d e n t i f i e d by d i s t i n c t i v e pelage colouration, occupied the same home range f o r periods ranging from 1.0 to 2.5 years. Enduring s i t e f i d e l i t y has been reported i n other coyote populations and appears to be a t r a i t common to other canid species. Camenzind (1978) found that some coyotes used the same home range during the four years of his study. Abies (1975) stated that there i s strong evidence that adult red foxes remain i n the same home range f o r l i f e . Also, adult wolves are known to use the same home range f o r considerable periods i f not l i f e (Haber 1977) . 5 6 Dispersal Dispersal i n mammals i s common at puberty and has been documented i n diverse groups such as voles (Krebs et a l . 1969), ungulates (Robinette 1966) and f e l i d s (Hornocker 1970). In canids, d i s p e r s a l i s a widely known phenomenon but quantitative data on the proportion of animals dispersing, time of d i s p e r s a l , and behaviour of dispersing canids are a v a i l a b l e f o r only one species, the red fox (see Storm et a l . 1976 for an excellent review). Information on coyote d i s p e r s a l i s l a r g e l y l i m i t e d to the distance t r a v e l l e d by each sex and the timing of d i s p e r s a l (Garlough 1940, Robinson and Cummings 1951, Knowlton 1972, Chesness and Bremicker 1974, N e l l i s and Keith 1976). In the present study, d i s p e r s a l information i s a v a i l a b l e for 12 coyotes, 11 of which were radio tagged and one which was ear tagged. I defined d i s p e r s a l as movement from one home range to another l o c a l i t y r e s u l t i n g i n a non-asymptotic r e l a t i o n s h i p between area u t i l i z e d and time. Dispersal ended when an asymptotic r e l a t i o n s h i p between area u t i l i z e d and time was established. Seven males and f i v e females dispersed (Table V I I I ) . This represents 46.2 percent of males and 38.5 percent of females radio tagged. Of the eight subadults (6 to 12 months) radio tagged and known to have been born on the study area, f i v e dispersed during t h e i r f i r s t winter. Two coyotes dispersed as y e a r l i n g s . A t h i r d y e a r l i n g (C50) was apparently dispersing when captured i n A p r i l and may have begun d i s p e r s a l as a subadult. Four adults ranging i n age from 25 months to 50 months changed home range during the study. Table VIIlJ Time of d i s p e r s a l and distance t r a v e l l e d by 12 coyotes of various-ages. Animal Sex Age (months) Date of v . i Distance d i s p e r s a l (km) t r a v e l l e d C l M 7 October 3 17.0 RRW M 11 March 17.3 C7 F 8 November 10.9 C17 F 7. October 11 49.2 C22 F 11 March 2 94.1 C20 M 22 February 19 16.6 C50 b M 13 NA C 64.0 C26 F 23 March 42.2 C12 M 30 August 15 3.4 C14 M 50 NA 8.5 C16 M 25 May 18 3.9 C15 F 43 October 25 7.1 Distance from point of capture to l a s t . r e l o c a t i o n . 'May have begun dispersing as a subadult. 1 'NA = not a v a i l a b l e . 58 Dispersal of subadults began i n early October and continued through March (Table V I I I ) . Two yearlings C20 and C26, l e f t t h e i r n a t a l ranges i n February and March r e s p e c t i v e l y . Of the three adults for which time of d i s p e r s a l was known, 2 l e f t i n the f a l l and one l e f t i n the spring. Eight of 10 coyotes dispersed between October and March. These data agree with the more extensive r e s u l t s of Storm (1965), Abies (1969) and Storm et a l . (1976) on red foxes i n that winter d i s p e r s a l i s most common. However, two adult coyotes i n t h i s study dispersed i n summer, one i n May and one i n Augus t. The distance t r a v e l l e d by dispersing coyotes i s w e l l documented and the data presented here agree with what i s known (Robinson and Cummings 1951, Hawthorne 1971, Chesness and Bremicker 1974, Hib l e r 1976). Usually an operational distance i s given which the i n d i v i d u a l must exceed i n order for d i s p e r s a l to be considered. This i s necessary i n the absence of d e t a i l e d information on the behaviour of i n d i v i d u a l animals. However, d i s p e r s a l may involve r e l a t i v e l y short distances. For example, C12 was a member of the Athabasca Pack u n t i l l a t e summer of 1975 as determined by radio telemetry and observations from November 1974. On August 15, C12 moved across the r i v e r into the Maligne Pack t e r r i t o r y adjacent to i t s former range (Fig. 15). The male spent progressively more time i n the Maligne Range u n t i l by early September i t was c l e a r the animal had become resident i n t h i s new area which was contiguous with i t s former range. C12 was s t i l l present on the Maligne t e r r i t o r y 17 months l a t e r . Size of home range Home range sizes of nine male and 11 female coyotes are presented i n Table IX. I used the mean where more than one estimate of home range s i z e 59 Table IX;. Home range s i z e of male and female coyotes i n Jasper National Park., Home range s i z e (km2) Males Females. Summer Winter Summer Winter CIO 77.7 66.0, 66.5 C2 17.9 C12 11.5 C3 5.0 10.9 ,. , C13 14.8 14.8 C4 10.2, 4.3 5.3, 8.2 C16 '14.6 C6 13.5 11.4 C18 7.1 17.4 C7 16.2 C19 12.4 C l l 15.8, 11.1 15.3 C20 17.5, 15.5 12.4 ' C15 9.8 43.8 C23 14.1 C22 10.5 C25 20.7 C24 10.5 • C26 11.2 C27 11.8, 60 was a v a i l a b l e f o r a si n g l e animal i n the same season. In these cases, each estimate was from a d i f f e r e n t year. In summer, male home ranges varied from 2 2 7.1 to 77.7 km and i n winter from 12.4 to 66.5 km . There was no s i g n i f i c a n t d i f f e r e n c e ( t = 0.41, d.f. = 9, P >0.05) i n the mean s i z e of male home ranges i n summer (13.9 ± 1.38 km2, n = 8) and winter (14.9 ± 0.89 km2, n = 3). Data f o r male CIO were omitted from t h i s analysis since h i s home range was unusually large. Furthermore, to include home range estimates f o r CIO would have considerably biased between-sex comparisons. Female coyote home range s i z e did not d i f f e r s i g n i f i c a n t l y (Wilcoxon sign rank t e s t ; T = 26, n = 8, 2 2 P >0.05) between summer (10.3 ± 1.05 km , n = 8) and winter (16.3 ± 4.11 km , n = 8). There was no s i g n i f i c a n t d i f f e r e n c e i n the s i z e of male and female home ranges (t = 0.22, d.f. = 18, P >0.05). Male home ranges averaged 14.2 ± 1.05 km2 (n = 9) and females averaged 13.3 ± 2.19 km2 (n = 11). If the a v a i l a b i l i t y of resources i s constant i n time and space then home range should increase as a function of group s i z e . To examine the re l a t i o n s h i p between home range s i z e and group s i z e , I c o l l e c t e d data during the winters, 1974-75 and 1975-76. Where possible I used the average of two or more radio-tagged coyotes to estimate group home range s i z e . However, three group home rangeswere estimated from the d i s t r i b u t i o n of observations (n) alone: the Athabasca Pack, 1974-75 (n = 75); the Rocky River P a i r , 1975-76 (n = 31); and the Maligne Pack, 1975-76 (n = 48). To standardize the d i f f e r e n t home ranges, I subtracted the water area within the home 2 ranges of the Talbot Lake Pa i r , 1974-75, - 3.0 km ; Rocky River P a i r , 1975-76, 2 2 - 9.2 km ; Cin q u e f o i l Pack, 1975-76, - 3.8 km ; and Maligne Pack, both 2 winters, - 1.3 km . 61 In coyotes, a d i r e c t r e l a t i o n s h i p e x i s t s between home range s i z e and pack s i z e (Fig. 11). The regression c o e f f i c i e n t , b = 1.9, d i f f e r s s i g n i f i c a n t l y from zero (P <0.001). About 87 percent of the v a r i a t i o n i n home range s i z e of packs i s explained by v a r i a t i o n i n pack s i z e . This r e l a t i o n s h i p suggests that coyotes increase the s i z e of t h e i r home range as group s i z e increases. However, large home ranges could be more productive and, therefore, support a greater number of animals. To demonstrate that group s i z e i s a causal f a c t o r determining home range s i z e , i t i s necessary to compare home range s i z e i n d i f f e r e n t years within the same family whose s i z e i s known to have varied. I assume here, that the qua l i t y of the range remains constant from year to year. The Athabasca Pack provides the best i l l u s t r a t i o n of how home range s i z e varies with group s i z e . In 1974-75, a pack of seven coyotes occupied 2 an area of approximately 20 km . The pack was reduced to four animals f o r 2 most of 1975-76 and i t s home range decreased i n s i z e to 12.7 km . S i m i l a r l y , a reduction i n the s i z e of the Maligne Pack from s i x to f i v e animals was 2 correlated with a decrease i n home range s i z e from 17.9 km i n 1974-75 to 14.8 km2 i n 1975-76. D i s t r i b u t i o n of coyote home ranges The s p a t i a l r e l a t i o n s h i p s among coyotes i n the Athabasca River v a l l e y from 1974 to 1976 are presented i n F i g . 12 to 15. Home ranges of radio-c o l l a r e d coyotes are indicated by 95 percent convex polygons. Only asymptotic ranges are shown. Home ranges of non-radio-tagged i n d i v i d u a l s , of radio-tagged coyotes for which i n s u f f i c i e n t data were a v a i l a b l e to 62 F i g . 11. R e l a t i o n s h i p between home range s i z e and pack s i z e during the w i n t e r s , 1974-75 ( • ) and 1975-76 ( O ). Home range s i z e estimated from the d i s t r i b u t i o n of observations. ^Home range s i z e c o r r e c t e d f o r water area. 20 CM J 16 N \n ft c n c ^ 8 E o 0 • a 9 -- b © o a o a,b o CD o. b © y = r = 5.7 + 1.9X .93 i 1 n = 10 1 2 3 4 Group 5 size 6 7 64 F i g . 12. The s p a t i a l d i s t r i b u t i o n of resident coyotes on the study area i n the winter of 1974-75. See text for explanation of map. i 65 66 F i g . 13. The s p a t i a l d i s t r i b u t i o n of resident coyotes and active dens ( • ) on the study area i n the summer of 1975. 68 F i g . 14. The s p a t i a l d i s t r i b u t i o n of resident coyotes on the study area i n the winter of 1975-76. 70 F i g . 15. The s p a t i a l d i s t r i b u t i o n of resident coyotes and active dens ( • ) on the study area i n the summer of 1976. 71 72 determine an asymptotic range, and of radio-tagged animals whose transmitters had f a i l e d are represented by a l i n e segment. The length of the l i n e was determined as that length which included 95 percent of the observations (n) of an animal; where n >10. Where two or more coyotes are indicated by one l i n e segment, these animals shared a common home range ( i . e . they were re g u l a r l y observed together or si n g l y i n the same space). I have not plotted the radio locations or observations of coyotes on these figures to avoid confusion. However, I have given an example of plotted radio locations using the data of females C3 and C6 during the winter, 1974-75 (Fig. 16). In F i g . 16 the number above each symbol indicates the number of times each coyote was found at various l o c a t i o n s . Also shown on F i g . 12 to 15 i s the 1219 m (4000 f t ) contour d e l i m i t i n g the habitable v a l l e y land. As many home ranges appeared to be shared by several coyotes and as the general l o c a t i o n of these home ranges persisted despite changes i n the i d e n t i t y of coyotes using them, I gave each shared range a name. Thus, f o r example, coyotes sharing the area from Jasper townsite to about 6 km north of Jasper were c o l l e c t i v e l y known as the Athabasca group. As many as 8-9 coyotes shared t h i s home range, although the number varied from year to year. Groups of three or more coyotes sharing a home range were termed packs i f they s a t i s f i e d c e r t a i n c r i t e r i a (see S o c i a l Organization). In the winter, 1974-75 (Fig. 12) r e l a t i v e l y few coyotes were radio tagged. Nevertheless, information on these animals, coyotes accompanying them, and observations of other groups made i t possible to del i m i t the home range of seven of nine suspected resident groups. Infrequent observations during t h i s f i r s t winter suggested that a group of coyotes occupied the area 73 Fi g . 16. D i s t r i b u t i o n s of radio locations f o r adult females C3 ( • ) and C6 ( A ) during the winter, 1974-75. The number above each symbol indicates the number of times each coyote was at various locations. 74 a e 4 4 4 75 centered about the Snaring River, while another used a home range between the Snaring group and the Athabasca group to the south (Fig. 12). These groups consisted of four p a i r s , three groups of three, a group of s i x and a group of seven or eight. S i m i l a r l y , eight shared home ranges were i d e n t i f i e d during the winter of 1975-76 (Fig. 14) c o n s i s t i n g of two p a i r s , two groups of three, two groups of four, a group of f i v e and a group of s i x . In addition to groups which p a r t i t i o n e d t h e i r use of space so that l i t t l e overlap of ranges occurred, some coyotes occupied home ranges which overlapped one (C4, F i g . 12) or more (CIO, F i g . 12 to 14) group ranges, but were never observed with members of these groups. Female C5's movements were also centered i n the area used by C4, however, she died before her r e l a t i o n s h i p with C4 or the other seven coyotes using the area could be determined. During both winters, except for the area near the Snake Indian River which was not studied, coyote groups were resident i n a l l areas of the Athabasca Valley. Similarly, i n summer, shared home ranges, each representing a breeding group, completely used the habitable v a l l e y land (Fig. 13' and 15). Known na t a l dens or dens used by pups for any period during the summer within each range are also shown on F i g . 13 and 15. The number*of coyote groups was generally stable from season to season and year to year. However, two changes were documented, one i n the summer of 1975 and one i n l a t e winter, 1976. The f i r s t involved the death of female C3 and the subsequent d i s p e r s a l of her mate and death of her pups, leaving the Talbot range vacant. Approximately three weeks a f t e r C3's death on June 29, 1975, the Rocky River pair (C6 and STF) were observed 3 km within the vacant Talbot range. Unfortunately, female C6's radio f a i l e d on 76 June 25, 1976, so that d e t a i l s of the Rocky River p a i r ' s range extension are not known. Observations over the next several months c l e a r l y established that the greater portion of the Talbot range had been taken over by C6 and STF. Conclusive evidence of t h i s was obtained when I discovered that C6 gave b i r t h to her l i t t e r i n summer 1976 i n the same den used by C3 the summer before. The remaining part of the Talbot range was apparently claimed by the C i n q u e f o i l group to the immediate south. The second change i n group ranges involved the formation of a new breeding p a i r from within an e x i s t i n g pack. Formation of the Kinross P a i r (Fig. 15) began i n l a t e January, 1976, when adult female C4 spent increas-i n g l y less time with other members of the Athabasca Pack. This change i n land use i s evident i n F i g . 17 where the frequency d i s t r i b u t i o n s of locations of female C4 and female C l l (the alpha female of the Athabasca Pack) are depicted i n three dimensions. During the period October-December, 1975, the use of the home range by both females i s v i r t u a l l y i d e n t i c a l . Female C4 began to r e s t r i c t her movements to one end of the home range during the period January-March, 1976. On February 28, 1976, C4 was observed with an unknown male. Her behaviour toward the male suggested that a pair-bond had been established. During t h i s same period, C l l continued to use a l l parts of the range but spent less time i n the area used by the pair than she had from October to December, 1975. F i n a l l y , from A p r i l through July, 1976, C4 and her mate and C l l and her mate occupied quite separate home ranges with l i t t l e overlap. ': Each female produced a l i t t e r of pups i n 1976. The l o c a t i o n of the respective den s i t e s i s indicated on F i g . 17 by the larges t peak on the three-dimensional surface f o r the period A p r i l - J u l y , 1976. 77 Fi g . 17. Frequency d i s t r i b u t i o n s of radio locations of females C4 and C l l during the period October, 1975 to July, 1976. Data for each animal are plotted on the same coordinate space. 78 c n n=66 O c t o b e r - D e c e m b e r , 1975 C 4 79 Discussion Ecologists have found the comparative approach a u s e f u l t o o l i n understanding rather diverse phenomena. To be e f f e c t i v e t h i s method requires a measurement standard. The study of mammalian space-use s u f f e r s from the lack of such a standard. This r e s u l t s from a lack of agreement on the b i o l o g i c a l meaning of the estimates generated from e x i s t i n g methods, rather than a paucity of techniques with which to estimate home range s i z e . Methods used to estimate home range s i z e f a l l i nto two classes: (1) those which assume no underlying u t i l i z a t i o n d i s t r i b u t i o n or home range shape ( S t i c k e l 1954, Mohr and Stump 1966, Sanderson 1966), and (2) those which assume an e x p l i c i t a p r i o r i model for the form of the d i s t r i b u t i o n (see Van Winkle 1975 f o r review). The f i r s t c l a s s of methods, an example of which i s the minimum convex polygon used i n t h i s study, s u f f e r s p r i n c i p a l l y from a marked dependence on sample s i z e f o r s i z e estimation and from the i m p l i c i t assumption that p r o b a b i l i t y of l o c a t i n g an animal i s equal at a l l points i n space. The p r o b a b i l i t y models (class 2 above) are affected l e s s by sample s i z e , but they usually assume a c i r c u l a r normal (Hayne. 1949, Calhoun and Casby 1958) or b i v a r i a t e ( e l l i p t i c a l ) normal d i s t r i b u t i o n of l o c a t i o n records (Jennrich and Turner 1969, Mazurkiewicz 1969) about the geometric mean (but see Ford and Krumme i n press). These assumptions, though'they may be j u s t i f i e d f o r c e r t a i n species, can lead to serious misinterpretations when the assumed form of the p r o b a b i l i t y d i s t r i b u t i o n i s wrong. I chose the convex polygon method because of i t s graphic s i m p l i c i t y , i t s wide h i s t o r i c a l use and s t a t i s t i c a l s t a b i l i t y (Jennrich and Turner 1969). Further, the method tends to provide estimates which are intermediate i n 80 value between two other widely used techniques: the minimum area polygon and the 95 percent p r o b a b i l i t y e l l i p s e . Using minimum convex polygons (Mohr 1947), Chesness and Bremicker (1974, unpubl.) reported the mean home range s i z e of 17 adult female 2 2 coyotes as 16.1 km and of 10 adult males as 67.1 km i n an area character-ized by mixed aspen-fir f o r e s t interspersed with a g r i c u l t u r a l land. 2 Gipson (1972) found that adult male home ranges averaged 32.8 km (n = 5), 2 whereas, adult female ranges averaged 13.1 km (n = 3) i n a grassland-scrub oak area of western Arkansas. In both studies, adult males had much larger home ranges than adult females. By contrast, I found no differe n c e i n the home range s i z e of male and female coyotes. S i m i l a r l y , Hibler (1976), using a regression analysis of home range s i z e on rate of increase of area u t i l i z e d by coyotes, found no s i g n i f i c a n t d i f f e r e n c e between the s i z e of - 2 - 2 male (x = 17.8 km , n = 7) and female (x = 20.2 km , n = 6) home ranges. It i s d i f f i c u l t to explain why males should have larger home ranges than females i n some areas and not i n others. One p o s s i b i l i t y i s that male coyotes are larger and therefore require a larger area to meet t h e i r energy demands. In coyotes, adult males are usually heavier than adult females. Hawthorne (1971) found that males weighed an average of 12.2 percent more than females. Daniel (1973, c i t e d i n Bekoff 1977a) found that males were generally 18.7 percent heavier than females. Males averaged only 4.9 percent heavier than females i n Jasper. Recently, Harestad and Bunnell (in press) established an empirical r e l a t i o n s h i p between home range s i z e (H) and carnivore body weight (W) i n the form H = 0.022 W1,3. 81 Using Daniel's data above as a worst-case example, t h i s equation predicts that males should have a home range approximately 24 percent l a r g e r than females. However, i n both studies that reported sex differences i n the s i z e of home ranges, those of males averaged 149 to 319 percent larger than those of females. Thus, differences i n the energy requirements of males and females are not s u f f i c i e n t to explain the data. Another possible explanation of differences between systems i n which there are male-female differences i n home range s i z e and those where there are not, may be rela t e d to the type of mating system. In species e x h i b i t i n g monogamy and a strong p a i r bond throughout the year, males and females occupy the same s i z e home range, excepting the period of early maternal care of young during which the female's range should be smaller than the male. This appears to be the s i t u a t i o n of coyotes i n Jasper. On the other hand, f e l i d s are generally regarded as promiscuous breeders that do not e s t a b l i s h a p a i r bond between breeding adults (Kleiman and Eisenberg 1973). Thus, i n 2 bobcats (Lynx r u f u s ) , Bailey (1974) found that male home ranges (x = 42.1 km , n = 4) were 2.2 times larger than the average for eight female ranges 2 (19.3 km ). Also, male bobcats are about 30 percent larger than females (Banfield 1974). Here again the greater energy requirements of the male are not s u f f i c i e n t to account for such large home ranges. In promiscuous species males maximize reproductive success by mating with as many females as possible. Thus, the large home range of males l i k e l y increases the p r o b a b i l i t y of encountering receptive females. Promiscuous mating has not to my knowledge been reported i n coyotes, but polygamy has been reported i n red foxes (Burrow 1968) and k i t foxes, Vulpes macrotis (Egoscue 1962). I 82 can think of no other function f o r the large home ranges of males reported by Gipson (1972) and Chesness and Bremicker (1974, unpubl.). An observation-area curve was not- determined f o r coyotes by either Gipson (1972) or Chesness and Bremicker (1974, unpubl.). Hibler (1976) provided observation-area curves, but plotted the independent v a r i a b l e i n increments of 50 radio l o c a t i o n s . This coupled with a sampling schedule that involved two three-day periods per month and one l o c a t i o n per hour within each period, account for the s i g n i f i c a n t l y greater time (x = 6.4 months, n = 5) required to achieve an asymptotic observation-area curve than i n the present study (x = 1.5.months, n = 17). The observation-area curve i s a us e f u l index to compare space use by i n d i v i d u a l s of d i f f e r e n t sex, age, or populations. I t i s important that standard methods of sampling and analysis be adopted. Observation-area curves have the immediate advantage of pr e d i c t i n g the sampling e f f o r t required to define an animal's home range. More time may then be spent c o l l e c t i n g e c o l o g i c a l and behav-i o u r a l correlates of home range parameters. In coyotes, I have shown a d i r e c t r e l a t i o n s h i p between home range s i z e and group s i z e . Furthermore, a change in.the s i z e of an i n d i v i d u a l pack between years resulted i n a corresponding change i n home range s i z e . Using data from Mech and Frenzel (1971), Mech (1973) and Van Ballenberghe (1975), I found that winter home range s i z e also v a r i e s d i r e c t l y with wolf pack s i z e , over a range of 5 to 10 i n d i v i d u a l s . Recently, several studies have documented a causal r e l a t i o n s h i p between family s i z e and t e r r i t o r y s i z e i n group-breeding birds (Parry 1973, MacRoberts and MacRoberts 1976, Woolfenden and F i t z p a t r i c k 1978) . Woolfenden and F i t z p a t r i c k (1978) suggest 83 th i s r e l a t i o n s h i p allows c e r t a i n male helpers to acquire a part of t h e i r n a t a l range as a breeding t e r r i t o r y . Since males are much more active i n t e r r i t o r i a l defense than females, i n h e r i t i n g part of a f a m i l i a r t e r r i t o r y i s of obvious advantage. In support of t h e i r hypothesis the authors presented several case h i s t o r i e s . A s i m i l a r strategy has been documented by Rogers (1978, Northwest Regional W i l d l i f e Society Meetings) for young female black bears. In t h i s species, female cubs acquired part of t h e i r mother's t e r r i t o r y and eventually, replaced, her upon her death. The formation of the Kinross P a i r from within the Athabasca Pack i n 1975-76 may represent a s i m i l a r phenomenon. I t seems clear that the younger female (C4) acquired her breeding t e r r i t o r y from the C l l and her mate's home range. Although the genetic r e l a t i o n s h i p of C4 and C l l was unknown, i t i s possible that C4, a y e a r l i n g when captured i n November, 1974, was a daughter of the older female ( C l l was about 6 years of age i n A p r i l , 1975). I have represented coyote home ranges as well-defined, s t a t i c e n t i t i e s . Obviously, t h i s i s a gross s i m p l i f i c a t i o n . The s i z e , shape and d i s t r i b u t i o n of use of a home range vary as an animal responds to events on p h y s i o l o g i c a l , behavioural and e c o l o g i c a l time scales. This dynamic behaviour i s r e f l e c t e d to some extent i n multiple asymptotes of the observation-area curve. In some cases multiple asymptotes res u l t e d from an increase i n home range s i z e (e.g. C l l , C4, C12, C13), whereas i n other cases home range boundaries s h i f t e d , but the t o t a l area used was the same (e.g. C6, C l l ) . In summary, adult resident coyotes generally used the same home range fo r several years and most l i k e l y f o r l i f e , but occasionally adults moved and established a home range elsewhere i n the population. Shared home ranges 84 were common throughout the year, but there was l i t t l e overlap between neighbouring group ranges. F i n a l l y , home range s i z e v a r i e d d i r e c t l y w i t h coyote group s i z e . 85 7 SOCIAL ORGANIZATION In the previous chapter, I demonstrated that resident, adult coyotes generally l i m i t t h e i r movements to a we l l defined home range. Also, I showed that many i n d i v i d u a l s appear to u t i l i z e the same home range as one or more conspecifics of the same or opposite sex throughout the year. In t h i s chapter, I consider the s o c i a l organization of coyotes i n Jasper National Park. By s o c i a l organization I mean the "complex of behavioural c h a r a c t e r i s t i c s determining the mode of dispersion of a popula-t i o n and the i n t e r - i n d i v i d u a l encounters within i t " (Crook 1965: 182). Results A s o c i a l c l a s s i f i c a t i o n of coyotes i n the population Coyotes were c l a s s i f i e d as transients, s o l i t a r y residents, a member of a pair or pack based on (1) the degree of s i t e f i d e l i t y exhibited, (2) the si z e and (3) composition of groups i n which they were r e g u l a r l y observed, and (4) t h e i r gregariousness (Table X). An i n d i v i d u a l coyote may be expected to belong to several or a l l of these classes during i t s l i f e t i m e . Further, an i n d i v i d u a l may belong to any given class more than once. For example, a coyote becomes a transient when i t disperses from i t s natal range as a j u v e n i l e or from i t s natal or other home range as an adult. I t may then become a s o l i t a r y resident, having established a home range, u n t i l such time as i t p a i r s . I f a mate dies, a coyote may once again become a transient or s o l i t a r y resident u n t i l a s u i t a b l e replacement i s found. Table .'X.r; C l a s s i f i c a t i o n of coyotes based on s i t e f i d e l i t y , group s i z e , group composition, and gregariousness. Class S i t e f i d e l i t y Group s i z e Group C o m p o s i t i o n G r e g a r i o u s n e s s Transient S o l i t a r y resident Resident* p a i r Pack b absent high high high 3-8 generally YOY or YLY, a but va r i a b l e age and sex v a r i -able adult male and f emale adults, YLY and YOY low low high, but probably lowest i n the f a l l high, but probably lowest i n summer oo OS ^OY - Young of year, YLY - Yearling. bGroup s i z e and composition vary seasonally with the production of young. 87 Transients - The v a l l e y population i n l a t e winter (March) consisted of approximately 15 percent transient coyotes (Fig. 18). Transients by d e f i n i t i o n exhibited l i t t l e or no s i t e f i d e l i t y and were r a r e l y observed with other coyotes excepting at ungulate carcasses contesting f or food. Of the 12 captured coyotes c l a s s i f i e d as transients at one stage of t h e i r l i v e s , eight were less than two years of age and four were adults. Five coyotes c l a s s i f i e d as transients represented animals dispersing from t h e i r n a t a l home range during t h e i r f i r s t year. In these animals, d i s p e r s a l was marked by a rather sudden, usually overnight, movement of several kilometers beyond the l i m i t s of t h e i r n a t a l range. The mean s t r a i g h t - l i n e distance t r a v e l l e d during the f i r s t day of d i s p e r s a l f or four juveniles was 6.6 ± 2.21 km. Subsequently the area u t i l i z e d by these animals s t e a d i l y increased, s i t e f i d e l i t y having been abandoned. An i l l u s t r a t i o n of t h i s behaviour i s given i n F i g . 19. Female C22 was approxi-mately s i x months of age when captured. .She occupied the same home range (presumably her n a t a l range) from November, 1975 to February, 1976'. On March 2, 1976, she was located 12.9 km north of her home range. During the next three weeks C22's index of area u t i l i z e d t r i p l e d . She was l a s t located on the study area on March 23, 1976. Four days l a t e r she was observed 94.1 km south of Jasper at the Columbia I c e f i e l d s . She had t r a v e l l e d a ' s t r a i g h t - l i n e distance of over 100 km i n four days. Two coyotes r a d i o - c o l l a r e d were dispersing when captured. I obtained only seven radio locations of y e a r l i n g , male C50 from A p r i l 12 to 16, 1975. During t h i s five-day period C50 t r a v e l l e d 13 km along the v a l l e y . Several times during the winter 1975-76 and again on June 7, 1976, C50 was observed i n Mt. Robson P r o v i n c i a l Park about 64 km ( s t r a i g h t - l i n e distance) from i t s ( 88 Fi g . 18. The number of coyotes . in' each of four s o c i a l classes i n March, 1975 (open bars) and 1976 (shaded bars). Percent of t o t a l number of animals i s given above each bar. 89 30 £ c 20 rd «*— o - Q 10 E 0 62 55 20 13 u 10 ' 15 15 Pack Pair So l i ta ry Transient 90 F i g . 19. Observation-area curve of subadult female C22 from Novem-ber, 1975 to March, 1976. Every second radio l o c a t i o n i s p l o t t e d . 7 0 0 r 600 A A A A A 500 • Nov o Dec B Jan D Feb * Mar 400 h 300 2 0 0 H O O O O O O B O B B B Q D n D 1 0 0 h e © © 9 9 0 10 2 0 30 40 50 60 70 8 0 Number of radio locat ions 92 l a s t known l o c a t i o n on the study area. S i m i l a r l y , adult, male C14 e x h i b i t -ed no s i t e f i d e l i t y (Fig. 20) during the four-week period he was located on the study area. He had t r a v e l l e d a s t r a i g h t - l i n e distance of 14 km from h i s capture s i t e . I do not know whether the animal l e f t the study area or the radio malfunctioned, but C14 was never resighted. Three coyotes dispersed from one home range and established a new home range a f t e r a period as a transient. Yearling male C20 dispersed on February 19, 1976; he had occupied the same home range for at le a s t 14 months and most l i k e l y since b i r t h . A f t e r approximately four months as a transient, C20 established a range about 7 km from h i s former home range. Adult, male C12 was a transient f o r less than four weeks from the time he l e f t the Athabasca Pack u n t i l he joined the Maligne Pack whose t e r r i t o r y lay on the opposite side of the Athabasca River. Adult female C15 l e f t her home range on October 25, 1975. For the next eight months she u t i l i z e d a s t e a d i l y increasing area u n t i l she established a home range approximately 10 km from the edge of her former range. S o l i t a r y residents - A small number of coyotes (10 percent, F i g . 18) held a home range, but remained unpaired and were observed r a r e l y i n the company of con s p e c i f i c s . I c a l l e d these animals s o l i t a r y residents. Four r a d i o - c o l l a r e d coyotes were s o l i t a r y residents f or a l l or a part of the study: an adult male (CIO, 6 y r ) , a y e a r l i n g male (C18), a y e a r l i n g female (G4) and an adult female (C15, 3 y r ) . CIO was captured on November 13, 1974, at the same s i t e female C3 was taken several weeks e a r l i e r . Radio locations of these animals over a several week period suggested that they were a resident p a i r . However, as the winter progressed the two spent less time with one another u n t i l i n 93 F i g . 20. Observation-area curve of adult male C14 from A p r i l , 1975, to May, 1975. Every second radio l o c a t i o n i s plotted. I 9 4 700 r 600 • • 500 N CD I— *6 400 - 300 X Of TJ C 200 • • D B Apr D May 100 i i i i i 10 20 30 40 Number of radio locat ions 95 December, 1974, i t was c l e a r that another male had replaced CIO as C3's mate. During the remainder of the study CIO was a s o l i t a r y resident. Of 24 observations, CIO was i n the company of another coyote only twice. CIO did not mate i n 1975 or 1976, although an observation on February 26, 1976, suggested that he may have attempted to p a i r . At 12:19, CIO was observed i n t e r a c t i n g with the male of a resident p a i r . The female's behaviour and the attention paid her by both males suggested that she was i n estrus. For approximately 5 min, CIO aggressively placed himself between the male and h i s mate, thwarting a l l attempts by the male to approach the female. Once, CIO investigated the female's perine a l area, but her t a i l remained down, and then lunged at the male as he approached. F i n a l l y , the male eluded CIO long enough to approach the female. She immediately raised her t a i l . The male proceeded to s n i f f her anal area several times. Subsequently, the female cantered o f f , followed c l o s e l y by her mate and CIO. Upon overtaking h i s mate, the male mounted the female several times, l i c k e d her vulva and then mounted her twice more. Meanwhile, CIO stood nearby but did not attempt to i n t e r f e r e . At 12:31, as the p a i r moved o f f together, CIO l e f t the area. Over the next hour, I observed a d d i t i o n a l courtship and mating a c t i v i t i e s of the p a i r . Both yearlings appeared to be s o l i t a r y residents i n t h e i r n a t a l home range. Evidence f or t h i s i s p a r t i c u l a r l y strong i n the case of C18 since he was captured at f i v e months of age. He spent the winter of 1975-76 and the summer of 1976 u n t i l h i s death, i n h i s n a t a l range. During that period he was never observed with pack members (n = 27, when either C18 or another member of the family was observed). S i m i l a r l y , y e a r l i n g , female C4 was never observed with any of the seven member Athabasca Pack for a period of f i v e months during the winter 1974-75. Yet during t h i s period she occupied the same area that was heavily used by the pack. Adult, female C15 was a s o l i t a r y resident from July, 1976 u n t i l she 96 paired sometime during the winter of 1976-77 p r i o r to February. Several observations i n February, 1977, indicated that a pair-bond between C15 and a male had d e f i n i t e l y been established. Resident p a i r s - In coyotes, an adult male and female share the task of feeding and rearing the young. T y p i c a l l y , t h i s adult male-female cooperation and asso c i a t i o n p e r s i s t s throughout the year. In Jasper, approximately 16 percent of the winter population consisted of resident pairs (Fig. 18): four pairs i n March, 1975 and three i n March, 1976. Both members of the pair u t i l i z e d the same home range throughout the year (Table XI and F i g . 12 through 15). I was unsuccessful i n radio tagging the male and female of any p a i r . However, based on observations of four coyote p a i r s , I found no differ e n c e i n the mean length of the home range of males (4.7 ± 0.74 km) and females (4.9 ± 0.90 km). Within t h e i r home range pa i r s produced a s i n g l e l i t t e r per year with the exception of the Rocky River Pair where i n 1975, female C6 f a i l e d to produce a l i t t e r . Frequent sightings and radio locations suggest that i f conception occurred the l i t t e r was aborted or otherwise l o s t early during the gestation period. She and her same mate produced a l i t t e r of f i v e pups i n 1976. P r i o r to the b i r t h of a subsequent l i t t e r , d i s p e r s a l and/or mortality of subadults reduced the summer family group to the adult breeding p a i r . The pair-bond between adult males and females may p e r s i s t for many years. Female C l l and male BRU were paired f o r the three years of the study. Also, female BLB and male WST produced a l i t t e r i n each of the study's three years. Packs - An average of 58.5 percent of the resident population i n the Athabasca Valley was organized into packs ranging i n s i z e from 3-8 adults 9 7 Table 'XI. Home range length of the males and females of four r e s i d e n t p a i r s based on observations. Home range length (km) P a i r Males Females Rocky R i v e r 5.25(36) a 6.25(62) Talbot Lake 5.75(49) 5.75(51) C i n q u e f o i l 5.25(5) 5.25(6) Coin 2.5(9) 2.25(15) aNumber of observations given i n parentheses. 98 yearlings and non-dependent young i n winter (Fig. 18). I use the term pack to r e f e r to a group of wild canids e x h i b i t i n g the following c h a r a c t e r i s t i c s : 1) a regular a s s o c i a t i o n of the same three or more i n d i v i d u a l s i n such a c t i v i t i e s as foraging, feeding, t r a v e l l i n g and r e s t i n g , 2) organization of group members by roles and/or dominance r e l a t i o n s h i p s , 3) r e s t r i c t e d membership to such groups, that i s animals are not constantly j o i n i n g and leaving one group then another, and, 4) members of the pack are c l o s e l y r e l a t e d k i n ; t h i s point i s op t i o n a l . Although i t i s l i k e l y that condition four w i l l be met i n most species, i t i s not e s s e n t i a l f o r a pack as I have defined the term. I believe these c r i t e r i a are implied i n any discussion of the s o c i a l canids. In t h i s section, I present data on the c h a r a c t e r i s t i c s of coyote packs i n Jasper. A t o t a l of 2,138 coyotes were observed on 1,137 occasions from June, 1974 to February, 1977. Observations of 46 dependent young were omitted from the summer data as the parent-young group i s an obligate r e l a t i o n s h i p . Thus, i f three young and two adults were observed, a group of two was recorded. In winter, not only are coyotes more v i s i b l e against the snow, but they tend to t r a v e l on frozen lakes and r i v e r s making i t more l i k e l y that a l l animals t r a v e l l i n g together would be observed. 99 In summer, 77.0 percent.of 339 observations were of sing l e animals and 23.0 percent were of groups of two or more (Table XII). Groups of three to seven yearlings and adults were observed on 37 occasions during the summer months. Of 798 observations i n winter, 45.2 percent were of groups of two or more and 25.5 percent were of groups of three to nine animals. When the number of coyotes observed i n groups of d i f f e r e n t sizes i s considered, the s o c i a l nature of the coyote i n Jasper i s even more apparent (Table XIII). In summer, 45.5 percent of coyotes observed were i n the company of one or more conspecifics and 28.4 percent were i n groups of three to seven animals. Of 1,613 coyotes observed i n winter, 72.9 percent were i n groups of two or more and 53.4 percent were i n groups of three to nine coyotes. Coyotes occupying the same home range may avoid one another or they may be organized into a s o c i a l group. In spring, 1975, s i x coyotes occupying the Athabasca range were radio tagged (see F i g . 13). Two members of the pack were not radio tagged. Chance observations of t h i s group the previous winter suggested that they were members of the same pack. This was confirmed by telemetry data. Daily simultaneous radio locations of these coyotes showed that they r e g u l a r l y t r a v e l l e d as a group. F i g . 21 i s a three dimensional i l l u s t r a t i o n of t h i s s o c i a l i t y ; the frequency d i s t r i -bution of radio locations of these s i x coyotes i s given f o r the period A p r i l to September, 1975. With the exception of C16, the d i s t r i b u t i o n of radio locations i s s i m i l a r f o r a l l coyotes. Male C16 dispersed from the Athabasca range i n l a t e May. This i s shown as a se r i e s of small peaks leading to the r i g h t of the f i g u r e . Table XII;i The percentage of observations of coyotes i n groups of d i f f e r e n t s i z e s a i n summer and winter, 1974-1977. Season Number of observations Size of group Summer Winter 339 798 77.0 12.1 7.4 1.5 1.2 0 ...9 0 54.8 19.7 10.4 6.5 4.3 1.5 ,2.8 0 aBecause packs often s p l i t into smaller groups, these figures are not e n t i r e l y accurate. Also, i n summer, dependent young are omitted from the analysis, as the parent-young group i s an obligate r e l a t i o n s h i p . TabJLeJXMll? The percentage of coyotes observed i n groups of d i f f e r e n t s i z e s 3 i n summer and winter, 1974-1977. Number of Season coyotes Size of group -1 .2 .3 4 5 " 6 . .7 . . . . 8 9 Summer 479 54.5 17. .1 15. .7 4. ,2 4. .2 0 4.4 0 0 Winter 1613 27.1 19. ,5 15. ,4 12. .9 10. ,5 4.5 9.6 0 .6 Because packs often s p l i t into smaller groups, these figures are not e n t i r e l y accurate. Also, in-summer, dependent young are omitted from the a n a l y s i s , as the mother-young group i s an obligate r e l a t i o n s h i p . 102 F i g . 21. Frequency d i s t r i b u t i o n of radio locations of s i x members of the Athabasca Pack from A p r i l to September, 1975. Data f o r each animal are plo t t e d on the same coordinate space. 103 C 2 0 ^ 104 Composition and s t a b i l i t y of coyote packs Two packs, the Athabasca and Maligne, were studied i n s u f f i c i e n t d e t a i l to discuss the composition and s t a b i l i t y of associations among coyotes. These accounts are based on approximately 160 hr of observation of coyotes feeding at ungulate carcasses, on chance sightings and r a d i o -telemetry information. I have given considerable d e t a i l i n t h i s section. This i s valuable because less d e t a i l e d information of t h i s kind has only recently been reported for t h i s species (Camenzind 1978). Athabasca Pack - In July, 1974, the Athabasca Pack l i k e l y consisted of two adult males, an adult female, four yearlings and four pups (Table XIV). Only the adult p a i r and four pups were a c t u a l l y observed together i n July. Two of the four yearlings were females radio c o l l a r e d i n November, 1974, within the range of the pack. One of:these females (C5) died of unknown causes i n l a t e December, 1974, whereas the other (C4) maintained a small home range within that of the pack's t e r r i t o r y throughout the winter of 1974-75. In l a t e November, 1974, seven coyotes comprised the pack. Each of these animals could be i d e n t i f i e d by d i s t i n c t i v e pelage c h a r a c t e r i s t i c s . The members of t h i s group were observed t r a v e l l i n g , sleeping and feeding together 75 times from November, 1974 to A p r i l , 1975. A l l seven members of the pack were observed together on 22 or 29.3 percent of these observa-tions and three or more members were observed together on 54 occasions. In April-May, 1975, f i v e members of the pack were captured and radio 105 Table >XIy>. A summary of the s i z e and composition of Athabasca Pack from July, 1974 to February, 1977. . Season Pack s i z e Composition Summer 1974 6-11 Winter 1974-75 7-8 Summer 1975 5-9 Winter 1975-76 Summer 1976 2-5 BRU AdM3 also l i k e l y C5 Y L Y F C l l AdF C4 YLYF 4 YOY C16 YLYM C12 YLYM BLS AdM BRU AdM BLS AdM C l l AdF C13 YOYM C16 YLYM C20 YOYM C12 YLYM C4 YLYF - d e f i n i t e l y member . . . . . by early A p r i l C5 YLYF - dead, December BRU AdM BLS AdM - not seen a f t e r Sept. C l l AdF C12 AdM - dispersed, l a t e July C13 YLYM C16 AdM - dispersed, l a t e May C20 YLYM LBG YOYF - not seen a f t e r July C4 AdF BRU AdM C l l AdF BRU AdM C l l AdF 5 YOY C20 YLYM - dispersed, January C4 AdF - paired with male from outside the pack, Feb. C13 YLYM - k i l l e d by car, March Winter 1976-77 BRU AdM C l l AdF 3 YOY - best estimate from tracks Ad - Adult, YOY - Young of year, YLY - Yearling, M - Male, F - Female. \ 106 tagged. From t h i s information and behavioural observations, I determined that the pack consisted of two adult males (BRU, BLS), one adult female ( C l l ) , two 2-year old males (C16, C12), one y e a r l i n g female (C4) and two y e a r l i n g males (C13, C20); a t o t a l of eight i n d i v i d u a l s . Yearling female C4 was f i r s t observed with the pack i n early A p r i l , 1975. Several changes i n group composition and s i z e took place i n the summer of 1975 (Table XIV). In May, 2-year-old male C16 dispersed. The only pup to survive the den was l a s t observed with the pack i n l a t e July. C12, a 2-yr old male, dispersed i n l a t e July or early August. And, f i n a l l y , BLS the other adult male was l a s t seen with the pack i n August, 1975. This l e f t only f i v e coyotes, three males and two females, i n the Athabasca Pack. In l a t e January, 1976, the Athabasca Pack was reduced to four animals by the d i s p e r s a l of male C20. In early February, 1976, female C4 s p l i t from the pack by r e s t r i c t i n g her a c t i v i t y to the northern part of the pack's t e r r i t o r y (see Chapter 6 and F i g . 17 for d e t a i l s ) . In March, 1976, male C13 was k i l l e d by a car, reducing the pack to the breeding p a i r of BRU and C l l . Female C l l gave b i r t h to f i v e pups i n A p r i l , 1976. A l l f i v e pups were t r a v e l l i n g with BRU and C l l i n l a t e July when the study was terminated. On a subsequent t r i p to the study area i n February, 1977, I observed BRU and C l l and obtained i n d i r e c t evidence (snow tracking) of three other coyotes t r a v e l -l i n g with the p a i r . Maligne Pack - The Maligne Pack consisted of s i x to eight i n d i v i d u a l s i n July, 1974: one adult male (WHS), one adult female, four pups, and l i k e l y two yearlings of unknown sex (Table XV). On October 9, 1974, I radio- tagged the adult female (C2) of the pack. From October, 1974 through January; 1975 the pack consisted of C2, her mate1 (WHS) and four coyotes 107 Table XV. A summary of the s i z e and composition of the Maligne Pack from July, 1974 to February, 1977. Season Pack s i z e Composition Summer 1974 8 C2 AdF a i WHS AdM UNK YLY UNK YLY 4 YOY Winter 1974-75 5-6 WHS AdM UNK YLY 3 YOY Summer 1975 9-10 C19 AdM TES AdF b C12 AdM -UNK YLY UNK YLY 6 YOY Winter 1975-76 5-6 C19 AdMc C12 AdM TES AdF C26 YOYF SLS YOYM UNK YLY? Summer 1976 9 C19 AdM C12 AdM TES AdF C26 YLYF SLS YLYM • 4 YOY Winter 1977 5-7 C19 AdM (February) C12 AdM TES AdF . 2-3 YOY C2 AdF - died, Jan. C26 YLYF dispersed i n March aAd - Adult, YLY - Yearling, YOY - Young of year, M - Male, F - Female, UNK - not known well enough to always be c e r t a i n of i d e n t i f i c a t i o n . ^May have been y e a r l i n g or adult female. c WHS and C19 l i k e l y same coyote. 108 whose sex and age were'uncertain. In l a t e January, 1975, the pack was re-duced i n s i z e to f i v e animals as a r e s u l t of the drowning of adult, female I C2. Apparently she was quickly replaced with another mature female as a l i t t e r of s i x was born i n A p r i l , 1975. Af t e r C2's death there was a three-month period where I knew r e l a t i v e l y l i t t l e about the s i z e or movements of the pack. However, on May 3, 1975, the adult male (C19) was radio tagged. It i s not clear whether the adult male c a l l e d WHS was the same animal as C19, although t h i s seems l i k e l y . During the summer of 1975, nine or 10 coyotes consisting of adult, male C19; adult, female TES; one or two y e a r l i n g of unknown sex, and s i x pups occupied the Maligne range. In August, 1975, C12 of the Athabasca Pack l e f t h i s range and over a period of several weeks'joined the Maligne Pack as the alpha male. By October, 1975, only s i x coyotes were observed r e g u l a r l y together. These included two adult males (C19, C12), one adult female (TES), one subadult male (SLS) and one subadult female (C26). The s i x t h coyote, believed to be a y e a r l i n g of unknown sex, was occasionally observed but spent progressively iess time with the pack during the winter. I l a s t saw t h i s animal on March.12,."1976, and i t seems l i k e l y that the animal dispersed. The fate of the three marked pups of the 1975 l i t t e r of s i x was unknown. They were l a s t observed i n early September. v In A p r i l , 1976, adult female TES gave b i r t h to four. pups. These nine coyotes were observed together several times throughout the summer and a l l ... A were believed to be a l i v e i n l a t e July when the study ended (Table XV.) . On a subsequent v i s i t to the study area i n February, 1977, the pack consisted of two adult males (C19, C12), an adult female (TES), a y e a r l i n g 109 female (C26) and two or three subadults. In March, 1977, C26 dispersed. This reduced the pack to f i v e or s i x animals. In each pack, the same coyotes associated with one another for periods ranging from 10 months to two years. Pack s i z e and composition varied annually depending upon the number and s u r v i v a l of young and the d i s p e r s a l of subadults, yearlings and two-year-olds. The Athabasca Pack varied i n s i z e from 11 i n the summer of 1974 to two i n l a t e winter 1975-76. During the same period the Maligne Pack varied between 5 and 10 adults, yearlings and young. Dominance re l a t i o n s h i p s within packs I assessed dominance based on the r e l a t i v e frequency with which i n d i v i d u a l s displayed a c t i v e or passive submission toward c o n s p e c i f i c s . These behaviours and t h e i r r o l e i n the dominance r e l a t i o n s h i p s of s i m i l a r species of canids, Canis lupus and C_. aureus, are discussed by Schenkel (1967) and Wandry (1975) re s p e c t i v e l y . A coyote which frequently exhibited submission to another coyote was considered to be subordinate to that animal. Behavioural observations were c o l l e c t e d p r i m a r i l y i n winter and thus, the r e l a t i o n s h i p s described should not be extrapolated to other seasons without due caution. I observed the Athabasca Pack f o r a t o t a l of 68 hr during the winters of 1974-75 and 1975-76. In 1974-75, the pack consisted of seven coyotes, s i x males and an adult female ( F i g . 22a). Adult male BRU was the dominant (alpha) animal. BRU was noticeably larger than male C13 and C20 each of which weighed 14.1 kg. The adult female ( C l l ) appeared to be the second ranking i n d i v i d u a l . I t was d i f f i c u l t to determine her rank because she 110 F i g . 22. Dominance re l a t i o n s h i p s among members of the Athabasca Pack i n winters, 1974-75 (A) and 1975-76 (B) and the Maligne Pack i n winter, 1975-76 (C). An arrow indicates that one coyote dominated another, whereas a dotted l i n e indicates that animals were of equal rank. I l l 112 displayed agonistic behaviour infrequently to animals other than BRU. Among males C12, C13, and C20 no cl e a r dominance could be determined. A l l three appeared to be of equal rank r e l a t i v e to one another and to other members of the pack. Yearling, male C16 was c l e a r l y the lowest ranking member of the pack behaving submissively i n agonistic encounters with other pack members. F i n a l l y , BLS, believed to be an adult male, r a r e l y interacted a g o n i s t i c a l l y with other pack members a f t e r a rather serious f i g h t with BRU i n January, 1975. Although' c l e a r l y subordinate to BRU h i s rank r e l a t i v e to other i n d i v i d u a l s was uncertain. However, on several occasions C16 displayed passive submission upon BLS's approach. The Athabasca Pack consisted of f i v e animals i n the winter of 1975-76 (Fig. 22b). BRU again.was the dominant coyote i n the pack. Males C13 and C20 appeared to be of equal rank as had been the case i n 1974-75. Female C4 also appeared to be equal i n rank to C13 and C20. A l l three of these animals co n s i s t e n t l y displayed submission to both BRU and female C l l . I observed the Maligne Pack f or 14 hr i n 1974-75 and 40 hr i n 1975-76. Consequently, comments on dominance w i l l be l i m i t e d to the winter of 1975-76 when f i v e coyotes comprised the pack (Fig. 22c). As was the case i n the Athabasca Pack, the alpha coyote was an adult male, C12. C19, also an adult, was the beta male and a j u v e n i l e SLS ranked t h i r d amongst the males. In the absence of the alpha male, C19 was dominant over the adult female TES.: However, when male C12 was present, C19 and TES r a r e l y interacted. Adult female TES was c l e a r l y dominant over both j u v e n i l e s . The r e l a t i o n s h i p between the juveniles was uncertain. Although they frequently interacted, the i n t e r a c t i o n generally took the form of play. In play, dominance r e l a -tions are often reversed and are generally u n r e l i a b l e (Bekoff' 1974). Too 113 few play bouts were observed to analyze the frequency of occurrence of play s o l i c i t i n g which Bekoff (1974) has shown to be correlated with dominance. D i v i s i o n of labour A c h a r a c t e r i s t i c of a l l highly evolved s o c i e t i e s i s a d i v i s i o n of labour among group members (Wilson 1975). In coyotes, d i v i s i o n of labour occurred with respect to ac t i v e defense of food and t e r r i t o r y , reproduction, scent marking, and group movements. I w i l l consider f i r s t , the con t r o l of pack movement. I observed members of the Athabasca Pack t r a v e l l i n g through t h e i r range 27 times. On 21 of these, the dominant male c o n t r o l l e d the speed and d i r e c t i o n of t r a v e l by leading the column of animals. In the remaining s i x instances c o n t r o l was exerted by the dominant male even though he was positioned elsewhere i n the column. This was evident i n the way that other members of the pack responded, to a change i n the behaviour of the dominant male. A s i m i l a r pattern was evident on 16 of 22 observations of t r a v e l by members of the Maligne Pack. However, the alpha male did not completely c o n t r o l the movements of other pack members. Individuals frequently l e f t the pack s i n g l y or i n pai r s p a r t i c u l a r l y when foraging f o r small food items. However, when the pack was t r a v e l l i n g as a uni t , the alpha male appeared to strongly influence where and when the pack t r a v e l l e d . The only exception to t h i s occurred i n the context of the breeding season. P a r t i c u l a r -l y when i n estrus the dominant female of a pack strongly influenced t r a v e l . Further evidence of the influence of the alpha male on pack movements comes from the behaviour of subordinate pack members a f t e r a period of r e s t . On several occasions, a l l members of the Athabasca Pack were r e s t i n g or 114 sleeping when I ar r i v e d . Several of the younger coyotes, e i t h e r s i n g l y or i n groups of two or three rose, stretched, yawned and then returned to t h e i r beds or engaged i n general a c t i v i t y within several hundred meters of other pack members. Over the .course of 30 min to an hour t h i s behaviour was repeated several times. Only when the alpha male rose and began to t r a v e l was a c t i v i t y transformed into organized movement. Before leaving, pack members usually e n c i r c l e d the alpha male and engaged i n a chorus v o c a l i z a -t i o n as they "nosed" one another and wagged t h e i r t a i l s . Throughout t h i s ceremony the dominant male maintained an a s s e r t i v e posture with h i s t a i l h o r i z o n t a l . D i v i s i o n of labour with regard to scent marking i n coyotes i s dealt with i n Chapter 8. However, several points w i l l be made here. F i r s t , my observations indicate that the dominant male frequently was the f i r s t and l a s t animal to mark a s i t e . Second, the dominant male scent marked approximately twice as often as other pack memb.ers. There i s circumstantial evidence that reproduction was l i m i t e d to one of several p o t e n t i a l breeding pa i r s i n the Maligne and Athabasca Pack. In the winter, 1974-75 the Athabasca Pack consisted of two adult males, an adult female and a y e a r l i n g female (C4) . However, only one l i t t e r of pups was produced and t h i s by the adult female. In 1975-76, female C4 paired with a male from outside the pack and produced a l i t t e r of f i v e young. As coyotes generally have t h e i r f i r s t estrus at 11 months of age (Gier 1968), at the age of 22 months ( i . e . winter 1974-75) C4 would have most c e r t a i n l y been p h y s i o l o g i c a l l y mature. 115 During the 1976 breeding season the Maligne Pack contained two females, an adult of unknown age (TES) and an 11-month subadult (C26). Two adult males and a subadult were also present. As was the case i n the Athabasca Pack only one l i t t e r and that by the adult female, was produced. That younger females i n these packs did not breed i n the presence of an older female need not imply a causal r e l a t i o n s h i p as several a l t e r n a t i v e explanations are possi b l e . I t i s possible both females became pregnant but l o s t t h e i r l i t t e r s e a rly i n gestation. The subadult female i n the Maligne Pack may not have ovulated. The proportion of 11-month-old females that ovulate i s highly v a r i a b l e i n other populations (Gier 1968, Kennelly pers. comm. c i t e d i n Knowlton 1972, Knudsen 1976) varying from 10 to 70 percent. However, the evidence suggests that further work i n t h i s regard i s needed. A d i v i s i o n of labour was evident i n t e r r i t o r i a l and food defense. When present, the dominant male took the i n i t i a t i v e against intruders i n 9 of 11 t e r r i t o r i a l confrontations and 8 of 9 instances of food defense. When the dominant male was not present another adult or high ranking y e a r l i n g ' generally played t h i s r o l e . Although usually successful, these males displayed less confidence than the alpha male. Genetic relatedness of pack members To determine .the kinship of members of coyote packs, I captured and i n d i v i d u a l l y marked l i t t e r s at dens or trapped j u v e n i l e s during t h e i r f i r s t summer. No young were marked i n 1974 as I was unable to reach the study area u n t i l a f t e r the young had abandoned t h e i r dens. In 1975, 13 young 116 representing four complete l i t t e r s and part of four others were ear tagged. In 1976, again no l i t t e r s were marked. Only one pup was found at the den used by the Athabasca Pack i n 1975. The pup was approximately 45 days of age when marked as determined by weight-age and body length-age regressions of Gier (1968). This pup was l a s t observed with the pack i n l a t e July and as i t was too young to disperse, i t must have died. Three of s i x pups of the Maligne Pack were ear tagged at approximately four to f i v e months of age. Although two young remained with the pack during the winter, 1975-76, neither was marked. The fate of the three marked i n d i v i d u a l s i s unknown. They were observed several times with the pack a f t e r t h e i r capture, but were l a s t seen i n September. Two pups of the Palisade Pack were marked at age 45 days. The l i t t e r of s i x was too young to ear tag when f i r s t captured at age seven days. Unfortunately, neither of these marked young was observed again. S i m i l a r l y , the complete l i t t e r of four of the Talbot Lake female were ear tagged at age 25 days. However, a f t e r her death a l l four young were believed to have died and her mate dispersed. Of the l i t t e r of two marked i n the C o l i n Pair's t e r r i t o r y i n 1975, one was k i l l e d by a car on December 23, 1975, approximately 0.5 km outside i t s n a t a l range and the second dispersed i n l a t e January, 1976. One three-month-old pup was marked i n the C i n q u e f o i l t e r r i t o r y i n 1975 and was not seen again. A second pup (C18) captured i n l a t e September, 1975, remained i n h i s natal range u n t i l h i s death i n June, 1976, almost 10 months l a t e r . 117 Unfortunately, these data provided l i t t l e information regarding the genetic relatedness o"f pack members. However, the age composition of three packs as determined by aging captured coyotes or as i n f e r r e d from the behaviour of coyotes suggests an extended family group. During the winter of 1975-76, the Maligne Pack consisted of three adults and two subadults from the 1975 l i t t e r of s i x . In A p r i l , 1976, the female subadult was radio tagged. Telemetry data indicated that she remained with the pack throughout the summer of 1976. She was s t i l l i n the Maligne range i n February, 1977, but dispersed i n March of that year. At the time of her death i n March, 1977, she was 23 months of age (age determined by counting cementum a n n u l i ) . S i m i l a r l y , the Athabasca Pack appeared to be an extended family.. During the winter, 1974-75, the pack consisted of adults, yearlings, and subadults (Table XIV). The Snaring Pack consisted of four coyotes during the winter, 1975-76. Two of these animals were smaller, engaged in play a c t i v i t i e s , and chased ravens (Corvus corax) considerably more than the larger animals. Therefore, I concluded the smaller animals were subadults. The subadults remained with the adult p a i r throughout the period of rearing the new l i t t e r and were s t i l l with the family i n July, 1976, when the study ended. The above evidence suggests that coyote packs are closed extended family groups. However, I recorded one instance where a coyote apparently unrelated to other members of a pack was accepted into the group. Male C12 was a member of the Athabasca Pack from the beginning of the study i n July, 1974, u n t i l he dispersed i n August, 1975. From September, 1975 u n t i l February, 1977, C12 held the p o s i t i o n of alpha male i n the Maligne Pack. 118 Land tenure Few mammals are t r u l y nomadic throughout t h e i r l i v e s , although a period of nomadic t r a v e l may often characterize the movements of dispersing subadults. More frequently, an i n d i v i d u a l establishes a home range (see Chapter 6) whereby i t becomes f a m i l i a r with an area within which i t conducts i t s normal d a i l y a c t i v i t i e s . In some species, i n d i v i d u a l s defend a l l or a part of t h e i r home range against c o n s p e c i f i c s . The defended portion of the home range i s known as a t e r r i t o r y . Defense may take the form of physi c a l confrontation with intruders or may include advertisement achieved through display, v o c a l i z a t i o n s , or o l f a c t o r y marking (Fretwell 1972). In t h i s section, I describe the t e r r i t o r i a l behaviour of resident p a i r s and packs. S o l i t a r y residents occupied w e l l defined home ranges, but I found no evidence that any part of the home range was defended. The nomadic behaviour of transient coyotes was discussed i n Chapter 6 and previously i n t h i s chapter. Eleven times a resident p a i r or pack confronted one or more intruders within t h e i r . t e r r i t o r y . In eight of 11 instances, intruders were vigorously chased to the t e r r i t o r i a l boundary. Fighting preceded three of these eight instances. Twice trespassers were followed to the edge of the te r r i t o r y , without a chase. Intruders were followed for 1.25 km i n one case and 50 m i n the other. F i n a l l y , on one occasion a v i s i t o r to an ungulate carcass near the edge of the Athabasca Pack's t e r r i t o r y was i n i t i a l l y chased 50 m by two members of the pack and subsequently followed for another 150 m. The en t i r e i n t e r a c t i o n was observed i n eight of nine 119 chases. Pursuit of intruders varied from 50 to 500 m with a mean of 218.8 m. In each instance, from two to f i v e residents were involved i n the chase. A t e r r i t o r i a l i n t e r a c t i o n , t y p i c a l of the r e s t , between the Maligne Pack and four intruders occurred on December 10, 1975. On December 8, 1975, I placed an ungulate carcass about.100 m east of the Athabasca River within the Maligne Pack's t e r r i t o r y . When I a r r i v e d at the carcass on December 10 at 08:19, three members of the Miette Pack and transient, female C17 were i n the v i c i n i t y of the food. The intruders had been at the carcass for about 90 min when I heard.the Maligne Pack v o c a l i z e about 1 km from the carcass. I observed no overt response i n any of the intruders at the carcass. Approximately a minute l a t e r , several coyotes probably other members of the Miette Pack vocalized f o r about 60 sec. Over the next 5 min coyotes on either side of the r i v e r exchanged v o c a l i z a t i o n s l a s t i n g on average about 20 sec. The coyotes at the carcass did not v o c a l i z e nor did they seem to respond to the c a l l i n g . At 10:15, four members of the Maligne Pack vocalized for 20 sec at a distance of no more than 300 m from the carcass. The intruders at the carcass continued to feed. F i n a l l y , at 10:30, the Maligne Pack rushed from the f o r e s t and quickly pursued the intruders. The intruders quickly retreated from the carcass, each i n a d i f f e r e n t d i r e c t i o n . The alpha male led the pursuit of the intruders. He and a subadult male quickly overtook two of the intruders and harassed them with b i t e s directed at the rump and shoulder areas. Simultaneously, the beta male and adult female chased one intruder about 200 m stopping only when the intruder had crossed the r i v e r and was 120 again i n i t s own t e r r i t o r y . A f t e r 30 sec of harassment the remaining two intruders bolted f o r the r i v e r . They were pursued 350 m by the alpha and subadult males of the Maligne Pack. Having chased o f f the trespassers, the Maligne Pack assembled and engaged i n a 15-sec chorus howl at the edge of the r i v e r . The alpha male scent marked and then the pack returned.to the ungulate carcass. At the carcass, each member of the pack scent marked either the food i t s e l f or objects nearby then slowly moved o f f into the woods i n the d i r e c t i o n from which they had come. At 10:39 the Maligne Pack again chorus howled for 15 to 20 sec. Three or four members of the Miette Pack immediately r e p l i e d . Associated with the above t e r r i t o r i a l i n t e r a c t i o n s involving p h y s i c a l confrontation was scent marking behaviour and v o c a l i z a t i o n s . Scent marking occurred i n s i x of the 11 i n t e r a c t i o n s , generally following the chase. Vocalizations accompanied only four of 11 i n t e r a c t i o n s . As i n the case of scent marking, v o c a l i z a t i o n s generally followed the chase (but see above). On 11 other occasions, resident pairs or packs harassed, attacked and/or chased intruders but did not pursue them to the edge of t h e i r t e r r i t o r y . However, i n two cases, the action taken by residents resulted i n the intruder leaving the resident's t e r r i t o r y . Nine of these 11 cases took place at ungulate carcasses within the resident t e r r i t o r y . In the other two food was not involved. In addition to d i r e c t p h y s i c a l confrontation, several authors have suggested that scent marking and v o c a l i z a t i o n s may serve to l i m i t the use of a resident's t e r r i t o r y by neighbours (Peters and Mech 1975, Lehner 1978). The use of scent marking by coyotes and the extent to which i t may be 121 regarded as t e r r i t o r i a l marking i s discussed i n Chapter 8. I c o l l e c t e d i n s u f f i c i e n t data to comment on the use of v o c a l i z a t i o n s i n t h i s regard. Discussion I suggested e a r l i e r that the s o c i a l organization of a large, mammalian carnivore depends to a large extent on the nature of i t s food resource. Thus, I w i l l defer much of the discussion of s o c i a l organization u n t i l I have considered coyote diets and foraging behaviour (Chapter 9). An e a s i l y obtained and relevant measure of s o c i a l i t y i s the percent frequency of observations of groups (excluding obligate parent-young a s s o c i a t i o n s ) . Although coyotes are usually observed as s i n g l e i n d i v i d u a l s , pairs and larger groups have been frequently reported. Ozoga and Harger (1966) observed pairs on 31.6 and larger groups on 7.4 percent of 228 winter sightings. Similar percentages of 30.0 and 6.0 were reported by Chesness and Bremicker (1974, unpubl.) i n 180 winter observations. In Jasper, approximately 20 percent of 798 winter observations were of pairs and 26 percent were of larger groups. Although comparable data are not presented, Camenzind (1978) r e g u l a r l y observed groups throughout the year suggesting that grouping i n t h i s population was s i m i l a r to that observed i n t h i s study, Percentages i n Jasper f a l l within the range observed for the highly s o c i a l wolf over a wide geographic area. Pairs of wolves comprised 14 to 28 and l a r g e r groups 22 to 48 percent of the 2,268 observations summarized by Mech (1970). Of the wolves observed i n these studies (n = 7, 477) 72 to 92 percent were i n groups. In Jasper, 72.9 percent of 1,613 coyotes observed i n winter were i n groups. Thus, there i s v a r i a t i o n i n the extent to which 122 coyotes form groups, and by t h i s measure alone, coyotes i n Jasper appear as s o c i a l as some wolf populations. A high degree of group-living i n coyotes has been reported only once previously (Camenzind 1978). Therefore, i t i s important to compare Camenzind's findings to the r e s u l t s of t h i s study. Working on the National Elk Refuge, Jackson Hole, Wyoming, Camenzind (1978) determined that the coyote population consisted of nomads (15 percent), resident pairs (24 percent), and resident packs of three to s i x coyotes (61 percent). Nomads were defined as coyotes that were r e g u l a r l y observed alone i n a home range which they did not defend. However, as home range was not defined rig o r o u s l y , i t i s l i k e l y that t h i s category includes my transient and s o l i t a r y resident classes. I f so, the Jasper population consisted of 25 percent transients and s o l i t a r y residents, 16 percent resident pairs and 59 percent packs. The s i m i l a r i t y i n organization of these two populations, e s p e c i a l l y with regard to the occurrence of packs, i s discussed further i n Chapter 10. Camenzind (1978) re f e r r e d to aggregations of winter migrants, nomadic coyotes, and residents as a separate organizational component of the population. These large transient groups (7 to 22 i n d i v i d u a l s ) were observed only i n winter near ungulate c a r r i o n . Camenzind found that these aggregations r a r e l y lasted more than 2 hr with an average duration of 38 min (n = 9 ) . Aggregations of t h i s type were common near ungulate carcasses i n Jasper, although the largest group observed was of 14 i n d i v i d u a l s . However, as these aggregations were so ephemeral, I do not see them as an organiza-t i o n a l segment of the population, but merely the r e s u l t of residents and non-residents contesting for food. On the basis of boundary scent marking, t e r r i t o r i a l disputes, and 123 v o c a l i z a t i o n s , Camenzind (1978) concluded that resident p a i r s as well as packs defended d i s t i n c t t e r r i t o r i e s . These data are consistent with the conclusions reached i n t h i s study. In Jasper, intruders were not always chased to t e r r i t o r i a l boundaries, e s p e c i a l l y when encountered near an ungulate carcass. This may seem to argue against t e r r i t o r i a l i t y . But, resident coyotes would expend considerably energy i n chasing intruders out of t h e i r t e r r i t o r i e s given the numbers of intruders attracted to a carcass. In these s i t u a t i o n s , coyotes probably b e n e f i t by only defending the food and not an area. At other times, defense of an area for prey searching i s f e a s i b l e . There seems l i t t l e doubt that the groups of coyotes found i n Jasper represent packs as I defined them e a r l i e r . Radio-telemetry data and observa-tions showed that members of these groups frequently t r a v e l l e d , s l e p t , foraged together (see Chapter 9) and displayed cooperative behaviour i n t e r r i t o r i a l defense. Agonistic i n t e r a c t i o n s between members of these groups r e f l e c t e d t h e i r s o c i a l status w i t h i n a dominance hierarchy. In coyote packs, a clear d i v i s i o n of labour existed i n the i n i t i a t i o n of t e r r i t o r i a l defense, scent marking, and the control of group t r a v e l . Also, I found circumstantial evidence of a r e s t r i c t i o n of reproduction to one of several p o t e n t i a l p a i r s , e s p e c i a l l y within the Athabasca Pack. Camenzind (1978) defined a pack as a group of coyotes that occupied and defended the same area, maintained dominance h i e r a r c h i e s , and often fed and denned together. Thus, h i s c r i t e r i a are s i m i l a r to those I used. He observed three packs from 1971 to 1974 with a d d i t i o n a l observations l i m i t e d to the denning period i n 1975 and 1976. During t h i s period, the three packs were comprised of three to s i x yearlings and adults i n l a t e winter. Camen-124 zind also reported circumstantial evidence of r e s t r i c t e d breeding. From 1971 to 1973, only one l i t t e r was produced although two females were present i n the Nowlin Creek Pack. In contrast, the Poverty F l a t Pack produced two l i t t e r s i n at least one of the 4 years of observations. I suspected that coyotes other than parents fed the pups, since they were r e g u l a r l y radio-located at the den. However, I was unable to f u r n i s h d i r e c t evidence of t h i s behaviour because of the d i f f i c u l t y i n observing coyotes i n the montane forests of Jasper. Helpers, defined as i n d i v i d u a l s which a s s i s t i n the feeding or caring of young other than t h e i r own (Skutch 1961), are common i n group-breeding birds (Brown 1974, Rowley 1974) and i n wolves (Haber 1977). Consequently, the observations of Camenzind (1978) and Ryden (1974) are p a r t i c u l a r l y important. Camenzind (p. 277) reported that "4 coyotes denned together and produced a t o t a l of 9 pups which were nursed by both females and fed solids:by both males with no apparent segregation of pups into i n d i v i d u a l s l i t t e r s . " Working i n t h i s same area of the National Elk Refuge, Ryden observed that two adults other than the parents fed the young and one of these (a female) r e g u l a r l y assumed a r o l e as alloparent. I conclude from these studies and my own observations that coyote and wolf packs, as described by Mech (1970), Fox (1975), Zimen (1976) and Haber (1977) are examples of the same type of s o c i a l organization. In each, i n d i v i d u a l s are organized i n a dominance hierarchy and show a d i v i s i o n of labour i n group t r a v e l , scent marking, t e r r i t o r i a l defense and care of young. Differences between wolf and coyote packs appear to be quantitative rather than q u a l i t a t i v e . For example, the s i z e of a wolf pack normally does not 125 exceed 15-20 i n d i v i d u a l s (Haber 1977) and 8-10 members i s most common (Mech 1970). In Jasper, coyote packs varied i n s i z e from 3 to 7 animals with an average of 4.4 (n = 8). S i m i l a r l y , i n Wyoming Camenzind (1978) found coyote packs averaged 4.7 i n d i v i d u a l s . Thus, i t i s l i k e l y that coyote packs are generally smaller than wolf packs. Coyote packs are probably less stable i n s i z e and composition than wolf packs. One reason f o r t h i s i s the behavioural ontogeny of these two species. Coyotes and wolves show behavioural differences as early as three to four weeks of age (Fox, 1969, Bekoff 1974, Bekoff, H i l l , and Mitton 1975). In c a p t i v i t y , coyotes t y p i c a l l y engage i n serious f i g h t s , the r e s u l t of which i s a clear-cut dominance hierarchy. This f i g h t i n g usually occurs between the t h i r d and f i f t h week of l i f e . S o c i a l play i s r a r e l y observed before domin-ance has been established. In contrast, wolves display l e s s a g o n i s t i c behaviour and more s o c i a l play early i n l i f e than coyotes. Although young wolves w i l l f i g h t o ccasionally, e s p e c i a l l y over food (Zimen 1975), they do not show early rank-related aggression. This has led several authors (Fox 1970, Bekoff 1977b) to suggest that the early development of agonistic behaviour i n coyotes impedes the development of strong f i l i a l bonds r e s u l t i n g i n the eventual d i s p e r s a l of young. A second reason coyote packs are l i k e l y less stable relates'!'> to the age at sexual maturity. Between 10 and 70 percent of 10-month-old female coyotes ovulate. In wolves, females normally do not ovulate u n t i l the age of 22 months. Thus, young wolves do not represent sexual r i v a l s u n t i l t h e i r second year or i n some cases t h e i r t h i r d year of l i f e . In coyotes, competi-t i o n among sexually mature o f f s p r i n g and parents l i k e l y p r e c i p i t a t e s d i s p e r s a l i n some instances. For example, male C20 was the second ranking 126 coyote i n the Athabasca Pack during the winter of 1975-76. At the onset of the breeding season, i n January, t h i s y e a r l i n g male suddenly disappeared from the pack. Three weeks l a t e r , I observed C20 i n the Athabasca Pack t e r r i t o r y . He had c l e a r l y been involved i n a serious f i g h t . His pelage was matted around the neck and hips and one ear appeared to be badly scarred where the ear tag had been. Male C20 was never observed with pack members a f t e r h i s i n i t i a l disappearance. His new home range was several kilometers to the south. Correlations of t h i s type do not provide evidence of cause and e f f e c t , but they are suggestive. To summarize, the s o c i a l organization of coyotes i n Jasper i s charac-t e r i z e d by a t e r r i t o r i a l mosaic of resident pa i r s and packs ranging from 3-7 in d i v i d u a l s i n winter. Superimposed upon t h i s mosaic are a number of s o l i t a r y resident home ranges, which overlap one or more resident t e r r i t o r -i e s , and the nomadic wanderings of dispersing coyotes ( t r a n s i e n t s ) . Coyote packs are composed of c l o s e l y r e l a t e d adults, yearlings, and subadults i n winter. This r e s u l t s from the delayed d i s p e r s a l of c e r t a i n subadults. For these i n d i v i d u a l s d i s p e r s a l usually occurs during t h e i r second winter. Male C12 and females C4 and C26 are examples of t h i s delayed d i s p e r s a l . Thus, there i s an annual turnover of subadults and yearlings with the breeding p a i r remaining as the nucleus of the pack. 127 8 SCENT MARKING Scent marking may be defined as the a p p l i c a t i o n of glandular secretions by an animal (also including urine and feces) to features of i t s environment. Few studies have investigated the frequency and d i s t r i b u t i o n of scent-marking behaviour of free-ranging mammals. However, these studies are e s s e n t i a l i f we are to evaluate the e c o l o g i c a l and behavioural s i g n i f i c a n c e of scent marking. Of the many functions ascribed to mammalian scent marking (see R a l l s 1971, Johnson 19 73) , > demarcation of a t e r r i t o r y has received more attention than e x i s t i n g data warrant. The •'.only evidence f o r t h i s hypothesis i n the w i l d i s i n the wild rabbit, Oryctolagus cuniculus (Mykytowycz 1968) and the wolf (Peters and Mech 1975). Within the Canidae, scent marking i s a widespread and w e l l documented phenomenon (Kleiman 19v66^ 19J.2) . The e c o l o g i c a l s i g n i f i c a n c e of t h i s behaviour has been the subject of much speculation. However, Peters and Mech's (1975) study was the f i r s t to document the d i s t r i b u t i o n and frequency of occurrence of marking and the response of animals to con s p e c i f i c scent marks for any cahid i n the w i l d . • L i t t l e i s known of the scent marking behaviour of free-ranging coyotes. In t h i s chapter, I describe the types and frequency of scent marking, the s p a t i a l and temporal d i s t r i b u t i o n of marking and the s t i m u l i that e l i c i t scent marking. F i n a l l y , I evaluate evidence of the t e r r i t o r i a l function of coyote scent marking and suggest topics f o r further study. 128 Methods Information was c o l l e c t e d p r i m a r i l y during the period of November through March, 1974-75 and 1975-76. I followed the tracks of one or more coyotes through undisturbed snow and recorded a l l signs of scent marking and eliminative a c t i v i t y . Scent marking was operationally defined as the o r i e n t a t i o n of urine and/or feces toward .particular objects i n the environ-ment (Kleiman 1966). Scent deposits without such o r i e n t a t i o n were consider-ed as eliminations. At each scent-mark l o c a t i o n , I recorded: number and type of scent deposits, number of coyotes present at the s i t e (within 2 m), type and numbers of objects marked and an estimate of the volume of urine present. The approximate volume of urine was estimated using a syringe loaded with coloured water (Peters and Mech 1975). If more than one urine or f e c a l deposit was recorded at one l o c a t i o n , I termed i t a multiple mark. These marks resulted from one animal marking several times or several animals marking the same object. Both occur, but I could not always d i s t i n g u i s h between them from tracks alone. Distances between scent-mark locations and t o t a l distance t r a v e l l e d by coyotes was measured with a pedometer. Results ' Types and frequency of occurrence of scent marks The analysis i s based on 1,047 scent marks of f i v e coyote packs and two mated pairs recorded along 731 coyote-km of snow tracks. Based on the 129 presence of urine, feces, and scratching, I recognized s i x types of scent marks: urine alone; urine and scratching; urine, feces, and scratching; feces alone; urine and feces; and feces and scratching. I assumed that each of these combinations could e l i c i t d i f f e r e n t responses from c o n s p e c i f i c s . Urine-alone scent marks occurred most frequently; 694 of 1,047 or 66.3 percent were of this type. To scent mark a coyote usually changed i t s r d i r e c t i o n of t r a v e l and approached an object upon which 0.5 to 2.0 cc of urine was sprinkled. In most cases, an average of 1.0 cc was sprinkled on a surface above the snow l i n e or on snowbanks and blocks of frozen snow. Peters and Mech (1975) suggested that depositing urine well above the ground may enhance d i s p e r s a l of odour by wind and minimize the chances that the mark may be covered with snow. Further, they noted that when placed in< the snow the urine l i k e l y constitutes a strong v i s u a l stimulus. Any surface appeared to be s u i t a b l e for the deposit of urine, however, the lower boughs of c o n i f e r s , tree trunks, small shrubs, t u f t s of grass, rocks, old feces, and ungulate bones were most frequently used. Urine and scratching scent marks comprised 28.7 percent of the 1,047 marks. Direct observation of coyotes marking indicated that a f t e r s p r i n k l i n g a small quantity of urine, coyotes generally took one or more steps away from the deposit before scratching the ground. Occasionally, both front and hind feet were used, but several alternate strokes of the hind limbs usually produced the scratch. The scratched area averaged 14.6 ± 1.4 cm wide by 61.8 ± 6.0 cm long (n = 50). More than one scratch was often produced for a given urine deposit. This behaviour appeared to be r e s t r i c t e d to the domin-ant male, but more data are needed to confirm t h i s . I 130 As well as representing a strong v i s u a l stimulus to conspecifics * (Peters and Mech 1975, Kruuk 1972, R a l l s 1971), scratching may also desposit secretions, of i n t e r d i g i t a l glands at the marking s i t e (Kruuk 1972). These glands are not known for c e r t a i n i n the coyote. However, coyotes do have eccrine sweat glands and apocrine (possibly odoriferous) sweat glands i n the foot pads (Sand, Coppinger and P h i l l i p s 1977). The 'debris loosened by scratching was usually directed away from the urine deposit and thus i s c l e a r l y not an attempt to cover the urine, as suggested by'"Young and Jackson (1951). The remaining four types of scent marks were r e l a t i v e l y uncommon. Scent marks containing urine,'feces, and scratching comprised 2.1 percent of the sample. Feces-alone scent marks were not recorded during t h i s study. Scent marks of urine and feces comprised 2.4 percent of the sample, whereas feces and scratching were recorded as marks only f i v e times. Generally urine and feces were oriented toward the same object, but the feces usually l a i d i n the snow. Often urine was sprinkled on top of the feces. D i s t r i b u t i o n of scent marks To examine the s p a t i a l d i s t r i b u t i o n of scent marks, each track was c l a s s i f i e d as l y i n g a) within a .25 km s t r i p along the perimeter of the t e r r i t o r y or b) i n the center. Data for the d i f f e r e n t packs were pooled a f t e r a Kruskal-Wallis one-way analysis of variance indicated no s i g n i f i c a n t differences i n the rate of scent marking among packs (H = 10.082, d.f. = 6, P >0.05) (Table XVI,). The mean number of a l l types of scent marks per coyote-km was s i g n i f i -cantly greater at the edge of the t e r r i t o r y (1.7 ± 0.16) than i n the center Table XVI. Summary of snow-tracking data from f i v e coyote packs and two mated pai r s i n Jasper National Park (scent marks within 20 m of k i l l or carcass excluded). Maligne Pack Atha-basca Pack Talbot Lake Pair Rocky River Pair P a l i s -ades Pack Snaring C i n q u e f o i l Pack Pack Sample s i z e 27 Distance tracked 426 (coyote-km) Total number of 592 marks 11 126 157 4 39 69 :.7 40 48 11 36 44 4 27 61 2 37 69 Marks/coyote-km 1.4C 1.2 1.8 1.2 1.2 2.3 1.9 \ No s i g n i f i c a n t difference i n rate of scent marking among packs; Kruskal-Wallis ANOVA H = 10.082, P >0.05. 132 (1.1 ± 0.12) (z = 5.715, n = 96, P <0.05) (Table XVlt). This increase at the edge of the t e r r i t o r y was brought about by increasing the number of scent marks per scent-mark l o c a t i o n and decreasing the distance between scent-mark s i t e s . At the edge, 70.3 ± 8.42 percent of scent marks were multiple marks compared to 50.5 ± 8.03 percent at the center of the t e r r i t o r y (Wilcoxon T = 0, n = 6, P <0.05). The mean distance between scent-mark s i t e s at the edge of the t e r r i t o r y (110.0 ± 6.00 m) was s i g n i f i c a n t l y less (t = 4.494, d.f. = 547, P <0.05) than i n the center (182.0 ± 13.00 m). Scent marks containing d i f f e r e n t information might be d i f f e r e n t i a l l y d i s t r i b u t e d throughout the t e r r i t o r y . However, the proportion of each scent-mark type d i d not d i f f e r s i g n i f i c a n t l y (Wilcoxon signed-ranks t e s t , n = 7, P >0.05 f o r each type of mark) between the edge and center of the t e r r i t o r y . A s i g n i f i c a n t increase i n the number of scratchings per coyote-km (Wilcoxon T = 1, n = 6, P <0.05) was observed at the edge of the t e r r i t o r y . On average, 0.54 ± 0.23 scratchings per coyote-km were found at the edge compared to 0.26 ± 0.12 i n the center of the t e r r i t o r y . Seidensticker et a l . (1973) reported that male .mountain l i o n s also scraped the ground more frequently near the edge of t h e i r t e r r i t o r i e s . We might expect the number of scent marks per animal to vary with the number, of animalsv,tracked. In coyotes t h i s does nojt appear to be the case. An analysis of variance indicated no change i n the rate of scent marking ei t h e r at the edge (H = 2.226, d.f. = 4, P >0.05) or i n the center of the t e r r i t o r y (H = 4.356, d.f. = 4, P >0.05) f o r pack sizes ranging from one to f i v e (Fig. 23). The independence of rate of marking and group s i z e suggests that a l l animals mark at approximately the same rate. A si n g l e coyote scent marked i 133 Table XVTT.' Comparison of rate of scent marking at the edge and center of coyote t e r r i t o r i e s . Sample No. coyote- No. of Marks/coyote-Location , s i z e km marks km Edge 51 406.7 707 1.74±0.163 S.E. a Center 45 324.9 340 1.05±0.123 S.E. d i f f e r e n c e between edge and center of t e r r i t o r y i s s i g n i f i c a n t , P <.001. 134 F i g . 2.3'. Mean rate of scent marking as a function of group s i z e at the edge (open bars) and i h the center (shaded bars) of a t e r r i t o r y . Sample s i z e and S.E. are indicated above each bar. 135 2.5r 15 E 2.0I i Oj O o 1.5 o O l (fi 1.0 2= 0.5 10 16 10 3 2 2 3 4 5 Number of coyotes 136 an average of 1.8 ± .31 times per km. Therefore, we can e a s i l y compute an expected value for d i f f e r e n t pack sizes assuming that a l l animals mark at the same rate. In F i g . 24 the upper regression l i n e indicates the expected r e l a t i o n s h i p between the number of marks per km and group s i z e . We see that the observed rates f a l l short of the predicted values above a pack s i z e of two. This suggests that the number of marks per km i s a function of age, sex, or s o c i a l status. Males and females of known age and r e l a t i v e dominance were observed to determine t h e i r rates of marking. Dominant males marked 12.7 ± 3.2 times per hr, whereas alpha females marked 6.6 ± 2.4 times per hr; a s i g n i f i c a n t d i f f e r e n c e (Wilcoxon T = 9, n = 18, P <0.05). I was'unable to determine the rate of marking of lower ranking or younger males and females. Alpha males also scratched s i g n i f i c a n t l y more often (33.0 ± 7.6 percent of marks were urine and scratching marks) than alpha females (18.9 ± 8.0 percent) (Wilcoxon T = 2, n = 15, P <0.05). Temporal d i s t r i b u t i o n of marking Although formerly thought to mark only during the breeding season (Kleiman 1966), I observed scent marking i n nine adult females throughout the year. These nine females were members of resident p a i r s and packs on the main study area. The data were c o l l e c t e d during chance observations and behavioural observations at ungulate carcasses. Table XVIII presents a summary of the number of dates each month that female scent marking was observed. These data should not be used to evaluate seasonal trends i n marking as unequal e f f o r t was applied throughout the year. The point to I 137 F i g . 24;'. Expected mean number of scent marks per km ( O ) assuming a l l coyotes mark at the same r a t e and mean observed r a t e s of marking ( # ) as a f u n c t i o n ' o f group s i z e . 138 T a b l e " X V I I I ^ Number of days each month that nine, adult females were observed scentVmarking. Female Jan Feb Mar Apr May Jun J u l Aug Sep . Oct Nov Dec C2 C3 1 1 4 2 1 2 1 1 C4 1 1 1 1 1 C6 3 2 2 2 2 ' 1 2 1 C l l 4 1 1 1 1 2 BLB 2 1 1 1 -ALI 2 1 1 TES 2 SAL 2 1 - < 1 1 The breeding season includes the months of January, February, and March. 140 note i s that females scent marked i n a l l months. The frequency of scent marking increases during the breeding season i n a number of canid species, including the coyote, i n c a p t i v i t y (Kleiman 1966). This has recently been documented i n free-ranging wolves by Peters and Mech (1975). Such an increase i n rate of marking was not evident i n Jasper (Fig. 25). I found no s i g n i f i c a n t d i f f e r e n c e ' i n the rate of scent marking among months for the period November to March either i n the center (H = 4.168, d.f. = 4, P >0.05) or at the edge of the t e r r i t o r y (H = 2.4367, d.f. = 4, P >0.05). However, more data are needed to confirm these findings since the sample sizes for November and December are small and no data are a v a i l a b l e beyond March. Stimuli e l i c i t i n g scent marking Captive studies suggest that one of the strongest s t i m u l i e l i c i t i n g canid scent marking i s the urine of an unfamiliar conspecific (Heimburger 1959, Kleiman 1966, F u l l e r and Fox 1969). Although i t i s clear that urine from neighbours i s a strong stimulus near t e r r i t o r i a l borders i n the f i e l d as well (Peters and Mech 1975, t h i s study), much of coyote scent marking cannot be explained i n t h i s way. Peters and Mech (1975) found that wolves responded strongly to t h e i r own scent marks even though other wolves had not been i n the area since the locations were l a s t marked. I noted s i m i l a r behaviour i n coyotes. On January 28, 1975 a section of road near the Maligne Pack's t e r r i t o r y was chosen to investigate remarking of scent marks. The .5 km section of road and a 10 m s t r i p oh e i t h e r side were checked each morning for 46 days. Three times during t h i s period many of the e x i s t i n g scent marks 141 F i g . 25. Relationship between number of scent marks per coyote-km and months of the year at the edge (•) and i n the center (O) of the t e r r i t o r y . Sample s i z e indicated above each symbol. 142 Nov Dec Jan Feb Mar 143 were remarked by the Maligne Pack even thought other coyotes were known not to have v i s i t e d the area. When I f i r s t began to monitor t h i s s ection of road only three scent marks, 100 to 150 m apart, were present. Over the next 46 days,ceight a d d i t i o n a l marking s i t e s were used by the pack or perhaps occasionally by neighbours (Fig. 26l). (As each new s i t e was marked, a numbered tag was placed on the nearest tree to define the location.) Coyotes used the road on 29 of the 46 days and marked at l e a s t one s i t e on 93 percent of these. Certain scent marks (1, 2, and 5) were remarked more often than others, and some were apparently not remarked (scent marks 8, 18, 24; F i g . 26). Inspection of these s i t e s indicated no obvious differences f i n p h y s i c a l char-a c t e r i s t i c s . Location of the mark along the road did not c o r r e l a t e with frequency marked. It seems that c e r t a i n s i t e s , also known to occur i n the center of the t e r r i t o r y , have s p e c i a l s i g n i f i c a n c e to coyotes: Two of the three frequently used s i t e s were d i f f e r e n t i n one obvious way. Of 18 scats deposited at scent-mark,sites along.the .5 km section of road, 14 were located at scent marks 1 and 2. Coyote urine and feces were c l e a r l y not the only s t i m u l i e l i c i t i n g scent marking i n coyotes. In over 80 percent of the scent marks recorded during snow-tracking, no previous mark (urine, feces, or evidence of scratching) was found at the s i t e . On s i x occasions marking behaviour was observed immediate-l y a f t e r a chase or encounter between residents and v i s i t o r s at a t e r r i t o r i a l boundary (see Chapter 7, Land tenure). 144 F i g . 26;. V i s i t a t i o n o f s c e n t marks by t h e M a l i g n e Pack a l o n g a .5 km s e c t i o n o f r o a d n e a r t h e edge o f t h e i r t e r r i t o r y . S i t e s marked ( • ) and s i t e s v i s i t e d but n o t marked ( O ) a r e i n d i c a t e d . Absence o f a symbol means t h a t a s c e n t mark was n e i t h e r marked n o r v i s i t e d t h a t day. 145 O t— f6 i — JQ CU 13 11 ; 9' 7 5 3J 1 2 7 2 5 2 3 21 19 17 15 13 11 9 30 281 O © O i o I o o I © I I I I o © X © o © o o I o °6 i o o o o I o I o o o I o o o 2 3 5 6 7 8 9 18 2 0 2 4 S c e n t - m a r k l o c a t i o n 146 Response to scent marks of neighbours I observed the responses of neighbouring coyotes to the urine of r e s i d -ent animals at ungulate carcasses located near t e r r i t o r i a l borders. Residents frequently scent marked on or near these carcasses. On one occa^.; sion, 22 separate urine deposits were recorded within 15 m of a carcass. These deposits were a l l less than 8 hr o l d . Thus, i n a very short time carcasses become o l f a c t o r y "hotspots" (Peters and Mech 1975) bearing resident scent. However, 22 times, trespassing resident coyotes showed no avoidance of these "hotspots" when they were alone at the carcass. A l l of these coyotes fed u n t i l s a t iated or displaced by other coyotes. Trespassing coyotes, i n marked contrast to residents, were never observed to scent mark or eliminate at or near a carcass. They also appeared anxious, responding strongly to the a c t i v i t y of ravens generally present at a carcass. Often the t a i l and hind quarters of these coyotes were lowered, also i n d i c a t i n g fear or anxiety. In the absence of an ungulate carcass, resident coyotes appeared to respect boundaries demarcated by scent marks. On three separate dates, I observed the reaction of a group of residents to fresh scent marks of another resident group at t e r r i t o r i a l boundaries. In each case the scent marks of the other residents were vigorously and systematically marked with urine and scratching scent marks. Both males and females scratched, but males more extensively. Subsequently, the residents returned toward the center of t h e i r own range. 147 Discussion Of the several means coyotes use to mark t h e i r environment, urine i s the most important. Feces are used, but there i s some evidence that t h e i r d i s t r i b u t i o n i s concentrated at c e r t a i n heavily reused s i t e s . Similar findings have been reported i n wolves by Peters and Mech (1975) and i n coyotes by Ozoga and Harger (1966), Gipson and Selander (1972), and Camen-zind (1978). In wolves, the alpha male and female do most of the scent marking (Peters and Mech 1975, Haber 1977). Consequently as pack s i z e increases the rate of scent marking decreases (Peters and Mech 1975). By contrast, the number of marks per coyote-km i s independent of coyote pack s i z e . As i n wolves, the dominant male of a pack.or p a i r of coyotes marks more frequently than lower ranking i n d i v i d u a l s . However, i n coyotes a l l pack members 9 months and older mark as often, or s l i g h t l y less often than the dominant female. This accounts for the increase i n marking rate with an increase i n pack s i z e (Fig. 24). It appears that d i v i s i o n of labour i n the use of scent i s less w e l l developed i n coyotes than i n wolves. Snowtracking and telemetry data suggest that coyotes t r a v e l over a large part of t h e i r t e r r i t o r y each day and that c e r t a i n routes e s p e c i a l l y at the edge of the t e r r i t o r y are used r e g u l a r l y . While t r a v e l l i n g , coyotes scent marked about every 150 m. Consequently, coyotes may encounter a recent scent mark approximately every 3 to 4 min, at t h e i r normal rate of t r a v e l . Under such conditions i t should be r e l a t i v e l y easy for coyotes to recognize the l i m i t s of t h e i r t e r r i t o r i e s (Peters and Mech 1975). This assumes that coyotes can i d e n t i f y group or i n d i v i d u a l odours. But i n d i v i d u a l recognition by urine or feces alone has not been demonstrated i n 148 canids. Mottus (1972) found no d i f f e r e n c e i n the time coyotes spent i n v e s t i -gating the urine of group members compared to non-group members. However, a l l the animals used i n t h i s test were of the same l i t t e r and may have shared chemical properties of urine common among l i t t e r mates. Kalmus (1955), however, demonstrated that dogs are capable of d i s t i n g u i s h i n g between the odour of human twins. I t seems l i k e l y that canids can d i s t i n g u i s h conspecif-i c s i n the same way (Ewer 1968) . Johnson (1973) suggested that scent marking as a means of t e r r i t o r i a l defense should be most frequent at the t e r r i t o r i a l boundary. Mykytowycz and Gambale (1969) have shown that dung h i l l s of the r a b b i t , Oryctolagus cunicu- l u s , are-.more numerous near a neighbouring colony, although marking s i t e s were d i s t r i b u t e d throughout the t e r r i t o r y . S i m i l a r l y , I found the density of coyote scent marks to be greatest at the edge of t h e i r t e r r i t o r y and lower i n the center. Coyotes appear to e s t a b l i s h an o l f a c t o r y "screen" around t h e i r t e r r i t o r y by reducing the distance between marking s i t e s and increasing the proportion of multiple marks. Peters and Mech: (1975) have demonstrated a s i m i l a r increase i n marking frequency by wolves at t e r r i t o r i a l borders. Thus, the edge of a t e r r i t o r y might be recognized by the rate at which scent marks are encountered;or, i f urine odour could be detected at considerable distances, an odour gradient might occur allowing coyotes to determine the l o c a t i o n of boundaries by the concentration of scent. However, the e f f e c t of wind would presumably l i m i t the effectiveness of t h i s method. Busnel (1963:74) reported the upper l i m i t of scent perception for wolves and dogs as 1500 to 2000 m. Increased scent marking at the edge of a t e r r i t o r y i s not s u f f i c i e n t cause to conclude that marking functions i n defense of a t e r r i t o r y . For , 149 example, captive animals mark i n s i t u a t i o n s producing fear, anxiety, or aggression (Ewer 1973). Thus, the frequency of scent marking may increase near the edge of an animal's t e r r i t o r y , where the p r o b a b i l i t y of encountering a neighbour i s greatest. In t e r r i t o r i a l maintenance, we must further show that neighbours "respect" scent marks demarcating a t e r r i t o r y by l i m i t i n g t h e i r use of the area beyond these marks. My data c o n f l i c t on t h i s point. Coyotes trespass t e r r i t o r i a l borders i n the presence of an ungulate caracss, but appear to respect scent marks when such a stimulus i s absent. Neighbours c l e a r l y benefit by trespassing under these conditions, as they obtain food otherwise unavailable. Under most conditions, coyote scent marking may be an e f f e c -t i v e t e r r i t o r i a l defense either alone or coupled with other communication systems. But, when a r i c h food source i s present near a t e r r i t o r y boundary, d i r e c t confrontation observed seven times during the study, i s necessary to prevent i n t r u s i o n . Scent marking i n the center of the t e r r i t o r y i s less l i k e l y to function i n t e r r i t o r i a l behaviour. Although the influence of having scent d i s t r i b u t e d over the e n t i r e • t e r r i t o r y on an i n d i v i d u a l having broken through the o l f a c -tory "screen" should not be underestimated. Peters and Mech (1975) suggested that scent marking i n the center of the range i s used as a means of o r i e n t -ing separated group members. Henry (1976) demonstrated that red foxes urine mark in e d i b l e food remnants throughout t h e i r range. The urine signals "no food present" when the s i t e s are re-investigated at some future time. Foxes spent le s s time at these locations and thus increased foraging e f f i c i e n c y . 150 Ungulate carcasses bearing resident urine were not avoided by v i s i t o r s . This i s consistent with the remark by Johnson (1973) that "there appear to be no observations of animals withdrawing a f t e r encountering an a l i e n scent mark". V i s i t o r s always appeared "nervous" and were much less e f f i c i e n t i n feeding than were occupants of a t e r r i t o r y (Chapter 9). This behaviour may r e s u l t from the coyote being out of i t s own range, but resident odour may, also be important. Similar observations i n the rabbit have been reported by Mykytowycz (1965,. 19681). Also, the absence of scent marking by trespassers at ungulate carcasses mights minimize the s t i m u l i e l i c i t i n g aggression i n residents. I believe that to gain further ins i g h t into the e c o l o g i c a l s i g n i f i c a n c e of coyote scent marking the following types of studies are required: 1) removal experiments, whereby the occupant of a t e r r i t o r y i s removed but i t s scent marks are a) a r t i f i c i a l l y maintained and b) allowed to become in a c t i v e , 2) a r t i f i c i a l extension of the scent marks of an animal into the range of another, and 3) analysis of urine and other scent bearing substances to i s o l a t e the b i o l o g i c a l l y a c t i v e chemicals. Recent work by Jorgenson e t , a l . (1978) and Alborne and Fox (1971) on the chemical constituents of red fox urine and anal gland secretion indicates t h i s to be a promising area of research. 151 9 DIETS, FEEDING AND FORAGING BEHAVIOUR If the nature of the food eaten i s an important determinant of predator s o c i a l organization, then we need a clear understanding of what foods are eaten and i n what r e l a t i v e proportion these foods occur i n the d i e t to under-stand the observed s o c i a l structure. Fortunately, t h i s type of information i s rather easy to obtain for most carnivores either by d i r e c t observation of what i s eaten or by c o l l e c t i n g feces throughout the year and reconstructing the d i e t from undigested prey remains. Consequently, more i s known of t h e i r food habits than any other aspect of coyote biology (see Bekoff 1977a for a recent review). Schoener (1971:392) stated that hypotheses on the adaptive s i g n i f i c a n c e of group foraging are of three types: "1. The group may hinder feeding, but there i s some overriding advantage to gregariousness. 2. Animals aggregate i n propor-t i o n to concentration of food, and group s i z e per se may be of neutral s e l e c t i v e e f f e c t . 3. Groups allow animals to forage more e f f i c i e n t l y . " In t h i s chapter I examine the di e t s and foraging behaviour of coyotes i n Jasper. Attention i s given to the type of prey eaten, and to the s i z e of prey items. I describe the feeding behaviour at ungulate carcasses and the influence of pack s i z e on diets to i l l u s t r a t e benefits that coyotes derive from group foraging. Methods To determine coyote d i e t s , I c o l l e c t e d known-aged feces along frequent-l y used roads and t r a i l s throughout the year and from a c t i v e dens i n early summer. When possible, I removed e x i s t i n g feces from these areas to allow accurate dating of future samples. I distinguished coyote feces from those of sympatric carnivores on the basis of s i z e and general appearance. Adult 152 coyote feces were 10 to 20 mm i n diameter. Wolf and mountain l i o n feces were greater than 20 mm i n diameter. Feces le s s than 10 mm i n diameter and c o l l e c t e d within a 50 m radius of an a c t i v e den were from coyote pups. Red i • • . fox and lynx occurred at such low d e n s i t i e s that any bias r e s u l t i n g from i n -cluding t h e i r feces was considered n e g l i g i b l e . In the laboratory, I s t e r i l i z e d feces at 110°C f o r 2 to 4 hr to k i l l the ova of Echinococcus granulosus. Samples were then moistened and separ-ated'. Seeds, mammal teeth, feathers and other items thought useful i n i d e n t i f y i n g food were saved. I i d e n t i f i e d most mammal h a i r macroscopically based on the following c h a r a c t e r i s t i c s : maximum diameter, length, and colouration of guard h a i r s . However, microscopic examination of c u t i c u l a r scales and the medulla was a l s o used. I followed the procedure of Spence (1963) i n making semi-permanent casts of h a i r . Generally, I mounted f i v e sample hairs to obtain a represent-ati v e p i c t u r e of the scale pattern. The h a i r medulla was examined using a whole-mount i n g l y c e r i n e . Unknown h a i r samples were compared to photomicro-graphs of reference h a i r c o l l e c t e d i n Jasper or from the Cowan Vertebrate Museum at the U n i v e r s i t y of B r i t i s h Columbia.(Bowen unpubl.) and from e l s e -where i n Western Canada (Richardson and Carbyn 1976). The r e l a t i v e importance ( i . e . c o n t r i b u t i o n of c a l o r i e s ) of small prey i s generally overestimated and o f ^ l a r g e prey underestimated when animal d i e t s are determined from feces. To p a r t i a l l y correct t h i s bias, I c l a s s i f i e d food remains i n feces as 1) a major item: 40 percent or more of f e c a l volume (v i s u a l estimate), 2) a minor item: 5 to 39 percent of f e c a l volume, and 3) a trace item: l e s s than 5 percent of f e c a l volume (Knowlton 1964). Only major items were used to evaluate d i e t s . Six percent of feces examined contained two major items. In these cases, each item was recorded as .5 occurrence to maintain the o r i g i n a l sample s i z e . 153 I placed prey Into one of four s i z e classes to estimate the r e l a t i v e occurrence of d i f f e r e n t s i z e prey i n the d i e t (Table XIX). The classes r e f l e c t an increasing tendency or requirement for cooperative foraging. This i s true i f prey i s captured or scavenged, since a group may better defend large prey (classes 3 and 4) from competitors than a si n g l e coyote. Thus, i n class 1, no cooperation i s necessary to capture prey and prey i s not shared, whereas i n class 4, cooperation i s generally required to capture prey and food i s always shared. Prey weights are from Soper (1970) and Banfield (1974). To investigate the foraging benefits of belonging to a pack, I observed coyotes feeding on ungulate carcasses. These carcasses were generally road-k i l l e d elk, mule deer, or mountain sheep that I moved to an area where coyotes would be undisturbed. I was c a r e f u l not to supply a large, p r e d i c t -able source of food to any one area. Ungulate carcasses were skinned, unless coyotes had started to feed before I found the food. In t h i s way I hoped to avoid b i a s i n g food habit data towards large prey. In t h i s a n a l y s i s , v i s i t o r s were transient coyotes or coyotes known to be outside t h e i r home range. Feeding, measured to the nearest minute, was defined as the time coyotes handled food or were within 1 m of the carcass. Results Diet T h i r t y d i f f e r e n t foods, c o n s i s t i n g of 18 mammals, four b i r d s , one f i s h , f i v e f r u i t s , and two insect species were recorded as major items i n 1,967 coyotes feces (Table XX). In addition, an unknown number of grasses (Gramineae) were recorded. Two a d d i t i o n a l species of mammal, a f i s h , and Table XIX"; C l a s s i f i c a t i o n of coyote foods by si z e (weight) . Average adult weight of each species given i n parentheses. Weight of food Foods included Class 1: <100 g Class 2: Class 3: Class 4: 100-1100 g 1.1-30.0 kg 30.0-450.0 kg No cooperation necessary for capture of prey, food not shared. Cooperation may increase, but not nec-essary f o r successful capture of prey, food may at times be shared. Cooperation may often increase success and may be necessary for-successful capture of some species, food often shared. Cooperation generally necessary for capture of prey and useful i n defense of such food, food regularly shared. Peromyscus (21.6), Microtus (37.8), Phenacomys (28.6), Zapus (23.7), Clethrionomys (24.3), Synaptomys (24.2), insects, f r u i t s Spermophilus (492), Neotoma (325), Tamiascuir.us (216), Ondatra (1100) Lepus (1.4); Marmota (5.9); Castor (20.0); Canis (13.2); Cervus3 c a l f (17.0); Odocoileus, fawn (3.8); .Ovis, lamb (3.3 ); f i s h , Prosopium and Catastomus (1.5); birds, Anatinae and Tetraonidae (1.2) Cervus (270.0),, Odocoileus (75.0), Ovis (102.0), Aloes (400.0), Ursus (150.0) U l Table XX. Percent frequency of occurrence of major items i n 1,967 coyotes feces from 1974 to 1977. , Food type Winter 1973-74 Summer 1974 Winter 1974-75 Summer 1975 Winter 1975-76 Summer 1976 Winter 1976-77 Mammals Ungulates Aloes aloes Cervus canadensis (ad) Cervus canadensis (young) Odocoileus hemipnus (ad) Odoooileus hemionus (young) ,: Ovis canadensis Lagomorphs and rodents Lepus americanus • Spermophilus columbianus Castor canadensis Ondatra zibethicus . Miorotus sp. Peromysous maniculatus Cleithrionomys gapperi Phenacomys intermedins Marmota galigata Tamiaeaiurus hudsoniaus Others Ursus americanus Unidentified mammals Birds Insects Phyllophaga sp. Melanoplus sp. Plants Shepherdia canadensis Rosa woodsii Gramineae Unidentified food 18 29 29 18 2 18 3 19 4 2 3 5 3 1 10 2 2 1 1 14 3 25 39 11 1 1 1 12 14 6 12 2 19 1 17 1 2 1 27 32 1 2 20 2 3 16 4 10 24 1 16 2 7 3 1 1 2 22 20 27 10 Sample size 17 349 357 434 370 400 40 dSpecies occurring <1 percent of the samples are not l i s t e d . ^ C o l l e c t i n g ended i n July before insects abundant. 156 three species of f r u i t occurred as minor or trace items. Although a wide range of foods were taken, mammals comprised 84.3 ± 3.53 percent of summer feces and 94.9 ± 1 . 8 7 percent of winter samples. The rest of the summer die t was 6.7 ± 4.06 percent ins e c t s , 4.0 ± 0.58 percent vegetation, and 1.3 ± 0.33 percent b i r d s . I was unable to i d e n t i f y the food remains i n 2.7 percent of summer feces and 2.5 percent of winter feces. In winter, mule deer, elk, and Microtus occurred i n 71.5 ± 3.13 percent of feces. These :same three species and Columbian ground s q u i r r e l s (Spermophilus columbianus) comprised 68.7 ± 5.24 percent of summer samples. With the exception of winter, 1973-74, snowshoe hares (Lepus americanus) were uncommon i n the d i e t (Table XX). Marked seasonal changes i n the r e l a t i v e frequency of occurrence of d i f f e r e n t prey i n the d i e t are shown i n F i g . 27. Small mammals increased i n the d i e t from July to a mid-winter peak of 34.57 percent and then f e l l r a p i d l y i n December-January to 2-6 percent. In March, small mammals again increased r a p i d l y peaking i n April-May at 25-44 percent of the die t and then dropping to a summer low of 8-10 percent frequency of occurrence. During both the winters, 1974-75 and 1975-76, ungulates ( a l l ages except summer young) occurred i n a maximum of 84 to 92 percent of coyote feces. In summer, ungulates comprised from 10 to 51 percent of the d i e t . Columbian ground s q u i r r e l s f i r s t emerge from hibernation i n the second or t h i r d week of A p r i l . In both 1975 and 1976, the r e l a t i v e frequency of ground s q u i r r e l s i n coyote feces rose sharply to 46.0 and 21.0 percent re s p e c t i v e l y i n June-July (Fig. 27). The frequency of ground s q u i r r e l s i n the diet f e l l markedly i n August r e f l e c t i n g the beginning of winter hiber-nation. Thus, coyotes responded very quickly to the appearance of an 157 F i g . 2 7. Seasonal v a r i a t i o n i n the percent frequency of Columbian ground s q u i r r e l s ( A ), mice ( • ) , young cervids ( A ), and other ungulates ( a l l species >6 mon of age; o ) i n coyote feces. 8 S I 159 abundant food supply i n summer. The r e l a t i v e frequency of mule deer fawns and elk calves i n summer feces mirrors the ground s q u i r r e l pattern, but reaches i t s maximum approxi-mately one month e a r l i e r (Fig. 27). Young cervids occurred i n 39.0 and 47.0 percent of coyote feces i n June, 1975, and July, 1976, r e s p e c t i v e l y . A locust, Melarioplus sp.,"occurred i n up to 23.0 percent of feces i n August and September. Its occurrence was strongly seasonal and was recorded only i n trace amounts outside these two months. The f r u i t of Shepherdia  canadensis and Rosa woodsii were taken i n small quantities i n 1.0 to 6.0 percent of feces i n August and September. Seasonal v a r i a t i o n i n the use of foods by coyotes has been well documented elsewhere ( F e r r e l et a l . 1953; F i c h t e r , Schildman and Sather 1955; Korschgen 1957; Kriowlton 1954; Gipson 1972; Meinzer, Ueckert and Fli n d e r s 1975; Brown 1977; Weaver 1977). In summer, the number of species i n the d i e t generally increases and average prey s i z e decreases. These observations are consistent with my information (see Table XX). In Jasper, seasonal v a r i a t i o n i n d i e t i s best i l l u s t r a t e d i n the use of Columbian ground s q u i r r e l s and young cervids (Fig. 27). Mice and adult ungulates were quickly replaced by ground s q u i r r e l s and young cervids as the dominant items i n the d i e t f o r a 2-month period i n June and July. The extent to which young cervids i n the d i e t represent coyote preda-t i o n rather than scavenging young k i l l e d by other agents, i s unknown i n Jasper. However, the sharpness of the peak i n occurrence suggests that young cervids are vulnerable to various mortality agents at or s h o r t l y a f t e r b i r t h , but that they quickly become less vulnerable. Salwasser (1972) and Cook et a l . (1971) came to s i m i l a r conclusions about mule deer i n 160 C a l i f o r n i a and w h i t e t a i l e d deer i n Texas r e s p e c t i v e l y . Cook et a l . (1971) provided evidence that s u b s t a n t i a l fawn predation by coyotes does occur. Over a 2-year period, they placed radio trans-mitters on 81 fawns 1 to 12 days o l d . Of the 58 deaths, 93 percent occur-red i n the f i r s t month of l i f e and 7 percent i n the following 30 days. Coyote, predation was responsible f o r 53 percent of the deaths. In twenty-two percent of the deaths the cause was undetermined, although coyotes had fed on the carcasses. The percent frequency of occurrence of d i f f e r e n t s i z e foods i n the adult coyote d i e t i s shown i n Table XXI. Throughout the year, c l a s s 1 prey comprised an average of 24.4 percent of the major items i n feces. In summer, the percentage of these foods i n the d i e t was generally higher than i n winter. An average of 15.3 percent of summer feces contained class 2 prey. In winter, class 2 foods occurred i n only 2.0 percent of feces. S i m i l a r l y , class 3 prey were more frequent i n the d i e t . i n summer (x =21.7 percent) than i n winter (x =5.0 percent). However, the reverse was true of cl a s s 4 prey ( i . e . adult ungulates). In summer, an average of 31.7 percent of •feces contained ungulate remains, whereas i n winter the f i g u r e was 66.5 percent. Table XXI also i n d i c a t e s that 5.3 and 2.0 percent of food remains i n summer and winter feces r e s p e c t i v e l y were u n i d e n t i f i e d . I also analyzed the stomach contents of 26 coyotes k i l l e d on the study area following the method of Korschgen (1969). Of these, two were empty Table XXI. Percent frequency of occurrence of d i f f e r e n t s i z e prey i n 1,463 adult coyote feces. - Summer Winter Food class 1974 1975 1976 X 1974 -75 1975-76 x;-1 32. 0 32.0 13. 0 25. ,7 22. 0 24.0 23. 0 2 6. 0 21.0 19. 0 •15. ,3 2. 0 2.0 2. 0 3 16. 0 18.0 31. 0 21. .7 . , 4. 0 6.0 5. 0 41. 0 22.0 32. 0 31. ,7 66. 0 67.0 - 66. 5 Un i d e n t i f i e d 5. 0 6.0 5. 0 5. .3 4. 0 0.0 2. 0 Sample s i z e 292 293 111 357 370 162 but the remainder contained an average of 300.6 ± 93.4 g of food (wet weight). The stomach of one female, k i l l e d i n summer, contained 1,594 g ;(3.5 lb:)aof fresh elk meat. A male, k i l l e d i n winter, had 1,568 g of elk meat i n i t s stomach. The stomach analysis also indicated that coyotes r e l y to a great ex-tent on ungulates f o r food. Thus, cl a s s 4 prey comprised 86.4 percent of food (by weight) i n the samples. Class 1 foods accounted f o r 9.9 percent of the d i e t , while classes 2 and 3 combined comprised 3.8 percent of the food i n stomach samples. Resident p a i r and pack d i e t s To v a l i d a t e my method of assigning the o r i g i n of feces to a group of coyotes based only on c o l l e c t i o n l o c a t i o n , I compared known feces of the Maligne Pack to a sample of feces (n = 50) c o l l e c t e d i n t h e i r range i n the same time period. The known sample (n = 57) was c o l l e c t e d while snow-tracking t h i s pack during the winter, 1975-76. Both samples agreed on the percent frequency of occurrence of the three prey species comprising approximately 90.0 percent of the food remains (Table XXII). Thus, t h i s method appears to accurately r e f l e c t the d i e t of a pack as indicated from snowtracking.samples. I found marked v a r i a b i l i t y i n the r e l a t i v e frequency of foods i n the feces of d i f f e r e n t p a i r s and-packs i n both summer and winter (Appendix I I I ) . This v a r i a b i l i t y was r e f l e c t e d also i n the r e l a t i v e proportion of d i f f e r e n t s i z e prey i n p a i r and pack d i e t s i n winter (Table XXIII) 'and. to_a lesser extent i n summer (Table<XXIV). Much of t h i s v a r i a b i l i t y between groups undoubtedly was caused by the patchy d i s t r i b u t i o n of prey such as the 163 Table XXII. A comparison of the percent occurrence of three prey species i n Maligne Pack feces to a sample of feces of unknown o r i g i n c o l l e c t e d i n the pack's t e r r i t o r y . Prey type Maligne Pack Unknown f e c a l feces (%) samples (%) Mierotus Cervus Odocoileus 4.0 32.0 53.0 7.0 32.0 50.0 Sample s i z e 57 50 Table JtXIII^, Percent frequency of occurrence of each food class i n the winter feces of coyote pairs and packs. 1974-75 1975-•76 Group name Food class Cin Tal Pal Mai Ath Roc Pal Cin . Sna Ath Mai 1 70 5 25 9 11 18 22 37 30 56 24 2 5 0 5 1 0 7 7 0 3 0 1 3 5 5 5 5 3 18 7 3 3 0 1 4 20 91^ 60 82 85 57 63 58 62 44 73 Uni d e n t i f i e d 0 0 5 3 1 0 0 0 3 0 1 Sample s i z e 20 21 20 139 76 28 27 38 37 25 180 Group s i z e 2 2 3 6 7 2 3 3 4 4 5 TabTe/ffiCiy.^ Percent frequency of occurrence of each food class i n the summer feces of coyote pairs and packs. 1974 1975 1976 Group name Food class Ath Sna ' -:. P al -./ Ath Sna Cin Tal Mai / Ath 1 30 69 30 20 J 44 36 35 60 8 2 7 4 12 29 5 13 5 17 20 3 8 5 12 5 10 17 17 4 49 4 53 22 44 46 35 32 40 18 23 Unid e n t i f i e d 1 2 2 1 5 . 1 2 1 0 Sample s i z e 24 92 25 21 20 23 23 92 50 Group s i z e a 4 2 3 6 2 2 2 4 2 Includes only yearlings and adults. 166 Columbian ground s q u i r r e l , mule deer, and elk. Unfortunately, time did not permit me to determine the r e l a t i v e d e n s i t i e s of these species on each coyote t e r r i t o r y i n summer. Thus, further analysis i s not possible. If cooperative behaviour increases the a b i l i t y of coyotes to capture large prey or locate and defend ungulate carcasses from competitors, then the proportion of class 4 foods i n the d i e t should be a function of pack s i z e . However, prey density and s i z e of area searched f o r prey also should influence the frequency of class 4 prey i n the d i e t . In summer, a regres-sion analysis indicated no s i g n i f i c a n t r e l a t i o n s h i p between the proportion of class 4 prey i n the d i e t and summer pack s i z e (see Table XXIV for data used i n a n a l y s i s ) . In winter, I estimated the r e l a t i v e abundance of ungulate species on each coyote-group t e r r i t o r y by the number of animals k i l l e d by vehicles along the 43 km of highway running through the study area. T r a f f i c volume and speed were constant along t h i s section of road. The highway passed through a l l t e r r i t o r i e s used i n the analysis except the Maligne. However, road counts and observations indicated that the density of ungulates i n the Maligne Pack's t e r r i t o r y was the same as the contiguous Athabasca Pack. Therefore, I used the same index value for each of these t e r r i t o r i e s . The number of elk, mule deer, and combined ungulates k i l l e d per coyote t e r r i -tory are given i n Table XXV. Elk density was more or le s s constant over the study area, however, mule deer density varied considerably. I used seven coyote groups (Table XXIII) and combined data from s i x s o l -i t a r y residents to regress the percentage of t o t a l ungulates i n the winter d i e t on winter pack s i z e and ungulate density. As i n the case of the summer food regressions, I used the arcsine transformation on percentage of food i n / 167 Table XXVA. Number of vehicular ungulate deaths per coyote-group t e r r i t o r y i n winter, 1974-75 and 1975-76 combined. Coyote group Number of deaths Mule deer Elk T o t a l ungulates Athabasca 13 7 20 Ci n q u e f o i l 2 5 18 Maligne 1 3 b 7 b 20 b Palisades 0 5 9 Rocky River 4 7 15 Snaring 3 5 10 Talbot Lake 0 4 Includes animals k i l l e d by t r a i n . Given same values as Athabasca Pack since highway did not pass through Maligne Pack's range; see text f or more complete statement. 168 the winter diet before performing the analysis (Sokal and Rohlf 1969). I assigned an average ungulate density index to the s o l i t a r y residents i n the regression. This seemed reasonable since s o l i t a r y residents generally had a larger home range and, therefore, l i k e l y experienced le s s v a r i a t i o n i n ungulate abundance than pa i r s or packs. A stepwise multiple regression of percent t o t a l ungulate on winter pack s i z e and ungulate density was s i g n i f i -cant for pack s i z e ( F 1 1 Q = 5.08, P <0.05) (Fig. 28). When ungulate-prey density was added to the model, the regression was not s i g n i f i c a n t . However, pack s i z e accounted f o r only a small amount of the v a r i a t i o n i n the dependent 2 v a r i a b l e (R = 0.36). C l e a r l y , other factors were important i n determining the amount of ungulate food i n the d i e t of d i f f e r e n t coyote groups. Because the t o t a l ungulate class contained animals of d i f f e r e n t s i z e , I p a r t i t i o n ungulates into the two species most common i n the d i e t , elk and mule deer. A multiple regression of the percentage of elk i n the winter d i e t on pack s i z e and elk density i n each coyote t e r r i t o r y was not s i g n i f i -cant ( F 0 0 = 0.93, P >0.05). Neither pack s i z e (Fig. 29) nor elk density (Table XXVI) were good predictors of the frequency of elk i n the feces of d i f f e r e n t groups. A multiple regression of the percentage of mule deer i n the winter d i e t on pack s i z e and mule deer density was highly s i g n i f i c a n t (F^ ^ = 46.8, P=0.001). The addition of mule deer density to the stepwise regression 2 increased R by only a f r a c t i o n of 1 percent, since pack s i z e and mule deer density were highly correlated, r = .82 (P <0.05). Pack s i z e alone accounted for 83.9 percent of the v a r i a t i o n i n the dependent v a r i a b l e ( Fig. 30).. The regression of mule deer density on the dependent v a r i a b l e was also highly s i g n i f i c a n t (F.. Q = 11.60, P=0.008). However, mule deer density explained 169 F i g . 2 8. Relationship between the percentage of ungulates i n winter feces and pack s i z e during 1974-75 ( • ) and 1975-76 ( O ). Data from s o l i t a r y residents ( • ) combined for the-two years. \ 170 100 80 - 60 O o- 40 20 -• - • ^"^ o o — o • a r c s i n J Y - 36.2 +LA x - • r =.58 n = 12 1 I i i i i i 1 2 3 4 5 6 7 Winter pack s i z e 171 F i g . 29. Relationship between the percentage of elk i n winter feces and pack s i z e during 1974-75 ( • ) and 1975-76 ( O ). 172 50 40 ~ 30 u i— CD ^ 20 10 -o o o • o • o • o • • • 1 1 1 i i 2 3 4 5 6 7 Winter pack s i ze 173 F i g . 30. Relationship between the percentage of mule deer i n winter feces and pack s i z e during 1974:-75 ( • ) and 1975-76' ("O') . 174 175 Table XXVI. Comparison of the p r o p o r t i o n of ungulates i n the wi n t e r d i e t of three coyote packs between 1974-^75 and 1975-76. Frequency of occurrence i n feces given i n parentheses. Athabasca Maligne P a l i s a d e s Pack Percent Percent Percent t o t a l Pack Year s i z e deer e l k ungulates 1974-75 7 46.1(35) 29.9(22) 80.3(61) 1975-76 4 20.0(5) 20.0(5) 44.0(11) t s 2.81 a 0.18 3.31 a 1974-75 6 56.1(78) 19.4(27) 82.0(114) 1975-76 5 39.4(71) 22.2(40) 73.3(132) t s 2.97 a -0.61 1.86 a 1974-75 3 10.0(2) 20.0(4) 55.0(11) 1975-76 3 4-18.5(5) 33.3(9) 63.0(17) t S -0.74 -1.03 -0.55 cl D i f f e r e n c e s between years s i g n i f i c a n t at P <0.05. 176 only 56.3 percent of the v a r i a t i o n i n the percentage of deer i n the winter feces of d i f f e r e n t coyote groups. That pack s i z e (2) i s a better estimator of the dependent v a r i a b l e (1) than mule deer density (3) i s shown also i n the p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s . The p a r t i a l c o r r e l a t i o n c o e f f i c i e n t s r12 3 ( t* i e c o r r e l a t i o n between va r i a b l e s 1 and 2 with 3 held constant) was .794, whereas the p a r t i a l c o r r e l a t i o n c o e f f i c i e n t r ^ 2 w a s "~-0006. As shown i n F i g . 11, pack size.(2) and home range s i z e (3) are highly correlated. Therefore, larger packs i n searching a larger area w i l l encounter more dead mule deer than smaller packs. This alone might account for the r e l a t i o n s h i p between percent mule deer i n the d i e t (1) and pack s i z e . There was a s i g n i f i c a n t regression of percent mule deer i n the d i e t on pack 2 1 s i z e and home range s i z e ( F 1 9 = 41.5, P=0.001, R =0.82). The p a r t i a l c o r r e l a t i o n c o e f f i c i e n t 3 = .906 was s i g n i f i c a n t (P=0.0001),' but r ^ 2 = -.360 was not (P=.307). Thus, pack s i z e was a good estimator of the amount of mule deer i n the d i e t when the e f f e c t of home range s i z e was removed, however the converse was not true. The e f f e c t of pack s i z e on the proportion of ungulates i n the winter di e t was analyzed further by comparing the same pack from year to year f o r packs whose s i z e had changed or had remained constant. Three packs were used i n t h i s a n a l y s i s : the Athabasca, Palisades, and Maligne. I used Sokal and Rohlf's (1969, p. 607) method for t e s t i n g the equality of two percentages. The Palisades Pack had three members i n the winters, 1974-75 and 1975-76. In both winters, the percentage of deer, elk, and combined ungulates i n the pack's d i e t was not s i g n i f i c a n t l y d i f f e r e n t (Table XXVI). 177 Two packs changed s i z e from one winter to the next. In 1974-75, there were seven animals i n the Athabasca Pack. In 1975-76, t h i s was reduced to 4 coyotes for most of the f e c a l c o l l e c t i o n period. Correspondingly, the percentage of mule deer i n the d i e t decreased s i g n i f i c a n t l y (t = 2.81, P <0.05) from 46.1 percent to 20.0 percent. S i m i l a r l y , the proportion of t o t a l ungulates declined from 80.3 percent i n 1974-75 to 44.0 percent i n 1975-76 (t = 3.31, P <0.05; Table XXVI). Note that the percentage of elk i n the d i e t was not s i g n i f i c a n t l y affected (t = 0.90, P >0.05). The Maligne Pack declined from s i x coyotes i n 1974-75 to f i v e i n 1975-76. I found.correlated changes i n the proportion of t o t a l ungulates and mule deer i n the d i e t (Table XXVI). The incidence of t o t a l ungulates decreased s i g n i f i c a n t l y (t = 1.86, P <0.05) from 82.0 percent i n 1974-75 to 73.3 percent i n 1975-76. Deer i n the d i e t declined from 56.1 to 39.4 percent; a s i g n i f i c a n t change (t = 2.97, P <0.05). The proportion of elk i n the pack's d i e t did not change from one year to the next (t = 0.61, P >0.05). Comparison of adult and pup d i e t s In c e r t a i n groups of animals, notably birds (Tinbergen 1960, Royama 1970, Hartwick 1973), the young are fed d i f f e r e n t foods than the parents themselves eat. Frequently, t h i s difference i s i n the s i z e of prey; adults generally consume smaller prey than they fed t h e i r young. To determine i f th i s might occur i n coyotes, I c o l l e c t e d pup feces from dens and adult samples for approximately the same period from May to July. I used the Wilcoxon Rank-Sign Test to compare the proportion of each prey-size class i n the d i e t of pups and adults. The r e s u l t s are given i n Table XXVII. I found that the proportion of ) Table XXVIP. Comparison of the r e l a t i v e frequency of each food s i z e class i n adult and pup feces ' ~~" ' ' f r o m M a y t o July, 1974 to 1976. Percent Percent Percent Percent food class 1 food class 2 food class 3 food class 4 Sample s i z e Pack Adult Pup Adult Pup Adult Pup Adult . Pup Adult Pup Rocky River '74 8.0 7.0 25.0 10.0 8.0 45.0 50.0 41.0 12 29 Maligne '75 57.0 46.0 17.0 0.0 4.0 5.0 13.0 31.0 50 13 Talbot Lake '75 33.0 14.0 2.0 7.0 17.0 7.0 39.0 72.0 23 14 Palisades '75 51.0 23.0 21.0 0.0 0.0 71.0 10.0 6.0 11 31 Miette '75 5.0 0.0 75.0 85.0 10.0 .12.0 10.0 3.0 10 35 Athabasca '76 8.0 5.0 20.0 18.0 49.0 58.0 23.0 19.0 50 84 Rocky River '76 12.0 7.0 20.0 4.0 16.0 59.0 54.0 29.0 13 52 Snaring '76 22.0 8.0 22.0 24.0 0.0 1.0 55.0 46.0 9 25 Mean 24.5 13.8 25.3 18.5 13.0 ' 32.3 31.8 30.9 S.E. 7.22 5.20 11.40 7.52 5.63 10.19 7.07 7.98 Wilcoxon Test T=0,n= =8,P<0.05 T=8.5,n=i 3,P>0.05 T=4,n= =8,P<0.05 T=14,n= =8,P>0.05 179 class 1 prey was s i g n i f i c a n t l y greater i n adult (24.8 percent) than i n pup die t s (13.8 percent) (T = 0, n = 8, P <0.05). The proportion of class 2 and class 4 prey items did not d i f f e r between pups and adults. However, the incidence of class 3 foods was s i g n i f i c a n t l y greater (T = 4, n = 8, P <0.05) i n pup feces (32.3 percent) than i n adult samples (13.0 percent). Thus, coyotes fed t h e i r pups larger foods i n the form of young cervids than they ate themselves between May and July. This may allow parents to spend les s time foraging and more time at the den, thereby reducing t h e i r energy expenditures and possibly the r i s k of pup loss through predation. Camenzind (1978) reported two instances of pup predation by neighbouring packs of coyotes. In each case, the k i l l e d pups were unattended at the den. Camenzind also observed f i v e cases of successful defense against intruders near active dens. I do not know i f pup predation may have occurred i n Jasper. Feeding behaviour at ungulate carcasses I observed coyotes feeding on 29 ungulate carcasses p r i m a r i l y during the winters, 1974-75 and 1975-76. Usually, I positioned carcasses near t e r r i t o r i a l boundaries to encourage i n t e r a c t i o n between groups, but carcasses were also placed at the center of t e r r i t o r i e s f o r comparison. Generally, I allowed a period of three of four weeks before I placed another carcass i n a pack's t e r r i t o r y to minimize the influence of supplemental feeding on the s i z e or dynamics of coyote groups. Carcasses were generally consumed i n three days or l e s s . Carcasses located near t e r r i t o r i a l borders were attended by residents from contiguous t e r r i t o r i e s , s o l i t a r y residents, and transient coyotes. Once, 14 coyotes from three packs were observed at a deer carcass. Carcasses 180 located i n the center of t e r r i t o r i e s were v i s i t e d only by the occupants of the t e r r i t o r y and occasionally by t r a n s i e n t s . However, transients r a r e l y fed long i n these circumstances. One advantage to i n d i v i d u a l s that belong to a pack might be i n the p r i o r i t y of access to large food resources. - To t e s t t h i s , I recorded the number of v i s i t s to a carcass that r e s u l t e d i n an animal a c t u a l l y feeding and the number that resulted i n no feeding i n various s o c i a l contexts. A v i s i t was recorded i f a coyote approached wi t h i n 100 m of the carcass. Only non-satiated coyotes were used.in the a n a l y s i s . • Coyotes were satiated i f : 1) an i n d i v i d u a l ( s ) alone at the carcass d i d not feed, 2) a coyote v i s i t e d a carcass but did not feed and allowed others to feed that arrived l a t e r , and 3) dominant coyotes did not feed while subordinate fed. I assumed that animals that v i s i t e d a carcass, beyond t h e i r home range were non-satiated. Non-satiated pack members always fed (n = 24) when they were alone at a carcass within t h e i r t e r r i t o r y (Table XXVIII). S i m i l a r l y , i f a s i n g l e member of a pack v i s i t e d a carcass w i t h i n i t s t e r r i t o r y , i t always fed (n = 16) when other coyotes were not present. The presence of v i s i t o r s at a carcass had no e f f e c t on the feeding behaviour of i n d i v i d u a l s of a pack (X 2 = 0.13, d.f. = 1, P>0.05; Table XXVIII). However, s i n g l e pack members fed l e s s frequently (x 2 = 3.86, d.f. = 1, P <0.05) i n the presence of v i s i t o r s than when alone at the carcass. Alone at a carcass, s i n g l e v i s i t o r s d i d not d i f f e r i n t h e i r feeding behaviour from s i n g l e residents; they fed on 20 of 22 occasions (Table XXVIII). Competition from other v i s i t o r s a lso had l i t t l e e f f e c t on the number of. v i s i t o r s that fed. However, the presence of s i n g l e residents or Table. XXVIII. The number of v i s i t s i n which non-satiated residents and v i s i t o r s fed upon ungulate carcasses i n d i f f e r e n t s o c i a l contexts. Number occasions Residents : >1 fed none fed 2 a X Resident Pack^ alone (PA) ' 24 0 Single resident alone (SRA)' 16 0 -Pack and v i s i t o r s present (P-V) 11 1 PA & P-V:0.13 Single resident and v i s i t o r 5 3 SRA & SR-V:3.86 present (SR-V) V i s i t o r s >1 fed none fed V i s i t o r s alone (SVA) 20 2 SRA & SVA:0.43 Other v i s i t o r s present (V-V) 12 1 SVA & V-V:0.23 V i s i t o r s and Pack present (V-P) 5 15 SVA & V-P:16.25 V i s i t o r s and s i n g l e resident (SV-SR) >3 5 SVA & SV-SR:6.61 Two-way test of independence with Yates correction f o r continuity, kpack considered present i f 2 members at the carcass. °X 2 of >3.84 i s s i g n i f i c a n t at P = 0.05 with l . d . f . 182 the resident pack c l e a r l y a f f e c t e d v i s i t o r feeding frequency. V i s i t o r s fed on only f i v e of 20 occasions when the resident pack was present; a s i g n i f i c a n t reduction i n feeding frequency compared to sing l e v i s i t o r s alone at a carcass (X2 = 16.25, d.f. = 1, P <6:66l). Also, the feeding frequency of sing l e v i s i t o r s was reduced i n the presence of a sing l e resident (x 2 = 6.61, d.f. = 1, P <0.01). Another measure of access to food i s the time spent feeding. A one-way analysis of variance showed that the mean feeding time of residents did not vary s i g n i f i c a n t l y among s o c i a l contexts (F^ = 0.794, P >0.05; F i g . 31). Residents fed somewhat i n excess of 20 min whether v i s i t o r s were present or not. Single v i s i t o r s fed for the same length of time as si n g l e residents when both were alone at a carcass (t = 0.55, d.f. = 40, P >0.05; F i g . 31). However, v i s i t o r feeding time was s i g n i f i c a n t l y decreased i n the presence of competitors, as indicated by a one-way analysis of variance (F^ - 10.42, P <0.01). I performed a Duncan's New Mu l t i p l e Range Test to determine which treatment means d i f f e r e d . In t h i s t e s t , treatments that are underscored with the same l i n e do not d i f f e r at P = 0.05. The r e s u l t s are indicated below (see F i g . 31 for legend): SVA V-V SV-SR P-V. The presence of residents and p a r t i c u l a r l y packs caused a marked decrease i n v i s i t o r feeding time from 24 min when alone to 4 min. Disturbance by other v i s i t o r s did not s i g n i f i c a n t l y a f f e c t v i s i t o r feeding time. 183 Fig . 31. Mean time spent feeding at ungulate carcasses by residents (A) and v i s i t o r s (B). Ninety-five percent condifence l i m i t s and sample s i z e are given above each mean. PA: Pack alone at carcass, SRA: Single resident alone at carcass, P-V: Pack and v i s i t o r ( s ) present, SR-V: Single resident and v i s i t o r ( s ) , SVA: Single v i s i t o r alone, V-V: Several v i s i t o r s present, SV-SR: Single v i s i t o r and single resident present. 184 10 30 r 5.8 21 20 | 10h cn c "•5 0 Q, P A S R A P - V S R - V B 3 2 » 3 0 r £ 20|-10h 2 6 42 S V A V - V V - P S V - S R 185 Residents not only fed on a greater proportion of t h e i r v i s i t s and for longer time, but also spent longer near the carcasses than v i s i t o r s . ; Coy-otes were observed at ungulate carcasses f o r 113 hr. During t h i s period, residents were present f o r 93.5 hr (82.6 percent), whereas v i s i t o r s were present for 68.7 hr (60.7 percent). V i s i t o r s were alone at carcasses f o r only 19.6 hr or 17.3 percent of the time. Unfortunately, I have i n s u f f i c i e n t data to determine the e f f e c t of pack s i z e on feeding behaviour at carcasses. Within packs, I could e s t a b l i s h no c l e a r feeding order. At. times a l l pack members (up to seven) e n c i r c l e d a carcass. A g o n i s t i c i n t e r a c t i o n was generally l i m i t e d to baring of the teeth which seemed to space animals at the food. Occasionally, a member of the pack might be chased s e v e r a l meters from the food, but i n v a r i a b l y i t quickly returned to feed. Ungulate predation I documented eight instances of ungulate predation by coyotes during the study: s i x mule deer and two Rocky Mountain sheep. Of the deer, two were adult females and four were fawns (two females, one male and one of unknown sex). As evidenced by the colour and firmness of the bone marrow i n the femur (Cheatum 1949), both adults were i n f a i r condition, although one of the females was rather gaunt i n appearance. Two of the fawns showed marked f a t depletion-, the other two had the bone marrow of a w e l l nourished animal. 186 An adult, female mountain sheep was k i l l e d by coyotes on February 8, 1975. The k i l l was reported to me by a warden days l a t e r . Unfortunately the bone marrow of t h i s animal was not inspected. The warden's account of the s i t e suggested that two or three coyotes had chased the sheep down a rather steep embankment and had pulled i t down as the sheep floundered at the base of the c l i f f . The second instance of sheep predation by coyotes involved a male lamb k i l l e d on December 11, 1975. This animal was i n good condition as judged by bone marrow c h a r a c t e r i s t i c s . In t h i s case,I was able to reconstruct the events that led to the k i l l with considerable d e t a i l . Two or three coyotes surprised a herd of sheep that were foraging approximately 75 m from escape t e r r a i n . The d i r e c t i o n of the coyote pursuit forced a part of the herd even further from the refuge of the nearby c l i f f s . A f t e r about 100 m of pursuit a si n g l e sheep (the k i l l e d lamb) either broke away or was forced from the herd by a coyote. Within 50 m a s i n g l e coyote had captured and k i l l e d the sheep apparently by suffocation. One of the adult female mule deer recorded as a coyote k i l l was a c t u a l l y k i l l e d by a park warden. Snowtracking data indicated that four coyotes (members of Maligne Pack) had chased and attacked the female i n f l i c t i n g serious wounds to the inner thigh and anal area (Fig. 32). Although,the deer apparently escaped by entering the r i v e r , i t was greatly weakened and would have died or been k i l l e d by the pack that night. At the time the deer was k i l l e d the Maligne Pack was approximately .5 km away. None of the k i l l s were observed d i r e c t l y , but tracks i n the snow allowed c e r t a i n inferences to be made. Generally, two to four coyotes xjere 187 F i e . 3 W o u n d s sustained by an adult female mule deer during a coyote predation attempt. 188 189 involved i n the pursuit and most l i k e l y i n the k i l l . Pursuit v a r i e d from 5 m when a deer fawn was apparently awakened to 400 m i n the case of an adult deer. The average of s i x chases was 189 ± 63 m. Five times predation attempts were observed d i r e c t l y or i n f e r r e d from tracks i n snow. On October 23, 1974, the alpha male of the Maligne Pack was observed chasing a group of f i v e mule deer (three adult females and two young). He had closed to within 5 m of one adult before I l o s t sight of the group. In July, 1974, I watched threeccoyotes of the Maligne"Pack chase an adult female mule deer approximately 200 m before I l o s t them i n the woods. In A p r i l , 1976, a s i n g l e adult male chased a group of three adult, female mule deer 100 m before the group disappeared into the woods. Twice i n March, 1976, chases were i n f e r r e d from tracks. In both cases, a s i n g l e mule deer had been chased by two or three coyotes approximately 100 m. Coyote predation i s probably more common than indicated above. In the only intensive attempt to locate k i l l s , I found four, a l l mule deer. A l l were located within the Maligne Pack t e r r i t o r y as a r e s u l t of 15 days of snowtracking over a 52-day period i n February-March, 1976. Mule deer appear to be the primary target of coyote predation i n Jasper. In a ddition to the cases reported above, 10 of 11 c o y o t e a k i l l s or suspected k i l l s (2 of 11) reported by wardens since 1966 were mule deer. The one exception was an adult, female mountain goat (Oreamnos americanus). 190 Discussion Five hypotheses have been proposed i n which group foraging i s considered to have increased foraging e f f i c i e n c y (Schoener 1971). The f i r s t two suggest that group foraging increases the a v a i l a b i l i t y of prey e i t h e r a c c i d e n t a l l y or purposely. Rand (1954) and Moynihan (1962) propose that group foraging may f l u s h more insects per b i r d than s o l i t a r y foraging. To some extent t h i s may occur when several coyotes are foraging f o r locusts, however, I believe t h i s i s generally unimportant i n t h i s species. In some carnivores t h i s may be s i g n i f i c a n t . For example, groups of k i l l e r whales, Orcinus orca, commonly herd dolphins, Delphinus sp. As i n d i v i d u a l whales rush into the middle of the school to feed, others remain c i r c l i n g . In t h i s manner almost the e n t i r e group of dolphins i s consumed (Brown and Norris 1956). A t h i r d and c l e a r l y more common benefit of group foraging i s an increase i n the s i z e of prey that can be captured (Schoener.1971, Kruuk 1972, 1975,.-\ Kleiman and Eisenberg 1973,; Alexander 1974, Wilson 1975). The d i r e c t r e l a t i o n s h i p between percent mule deer i n the winter d i e t and pack s i z e i n t h i s study suggests that coyotes may b e n e f i t i n t h i s way. However, there are several other possible explanations for t h i s r e l a t i o n s h i p . One p o s s i b i l -i t y i s that larger packs might search a larger area thereby encountering more deer that had died from causes other than coyote predation. Another explan-ation might be the larger packs defend and u t i l i z e a carcass more e f f e c t i v e l y than smaller groups or s i n g l e coyotes. F i n a l l y , large packs may l i v e i n areas of higher deer density, thus more dead deer would l i k e l y be a v a i l a b l e to large packs than small ones. 191 I found that the larges t packs did occur i n areas of highest deer density. However, when the e f f e c t of prey density was removed, pack s i z e was s t i l l a good estimator of d i e t , whereas prey density alone accounted fo r l i t t l e of the v a r i a t i o n i n d i e t . Also, the percentage of mule deer i n the winter d i e t varied within packs whose s i z e had changed and i n these cases i t i s u n l i k e l y that deer density had changed markedly. Thus, i t seems l i k e l y that the increase i n the proportion of deer i n the d i e t of larger packs i s brought about by an increase i n e f f i c i e n c y of some combination of capture, defense of food or searching f or food. The r e l a t i v e importance of these three factors i s d i f f i c u l t to deter-mine, however the u t i l i z a t i o n of elk by coyote packs provides some i n s i g h t . Elk predation by coyotes i s rare, although several cases have been reported (Cowan 1943, Weaver 1977). Consequently, the independence of amount of elk i n the d i e t from pack s i z e suggests that a p a i r of coyotes can locate ungulate carcasses as well as larger groups, since approximately the same number of elk were k i l l e d by v e h i c l e s i n each coyote t e r r i t o r y . In contrast to deer, the percentage of elk in.the winter d i e t did not vary within packs whose s i z e had changed from one year to the next. Because predation can be ruled out, the a b i l i t y of packs to locate elk carcasses appears independent of the number of searchers, above a minimum of two animals. Although smaller i n s i z e , i t i s l i k e l y that the a b i l i t y of packs to locate mule deer i s also independent of the number of searchers. We can also eliminate the s i z e of the area searched f o r prey, since when the e f f e c t of pack s i z e i s s t a t i s t i c a l l y removed, home range s i z e accounted for l i t t l e of the v a r i a t i o n i n the amount of deer i n the winter d i e t . Therefore, the 192 r e l a t i o n s h i p between the amount of deer i n the d i e t and pack s i z e l i k e l y i ndicates increased capture success or defense and u t i l i z a t i o n of a carcass. Increased predation success with an increase i n group s i z e has been reported i n other species. P a r t i c u l a r l y important are the observations on j a c k a l s , since they are s i m i l a r to coyotes i n s i z e , s o c i a l behaviour and ecology. Wyman (1967) found that the success of jackals i n k i l l i n g young Thomson's gazelle (Gazella thomsoni) increased from 16 percent when a s i n g l e j a c k a l was involved to 67 percent f o r a p a i r . Van Lawick and K f a n Lawick-Goodall (1970) found that groups of three to seven golden j a c k a l s , Canis  aureus, r e g u l a r l y hunted adult Thomson's gazelle. Schaller (1972) reported that the success of a s i n g l e l i o n (Panthera leo) i n k i l l i n g Thomson's gazelle, wildebeest (Connochaetes taurinus) and zebra (Equus b u r c h e l l i ) was 15 percent. Predation success of two l i o n s , however, increased to 29 percent, although l i t t l e improvement occurred with three or more l i o n s . S i m i l a r l y , Kruuk (1972) observed that a s i n g l e hyaena (Crocuta crocuta) was successful i n only 15 percent of i t s hunts of wildebeest calves, whereas two or more hyaenas k i l l e d wildebeest calves an average of 73 percent of the time. Several authors have observed that there i s a r e l a t i o n s h i p between.the si z e of prey and the number of hunters (Kruuk 1972, 1975, Schaller 1972, Eaton 1974). Thus, adult wildebeest are usually hunted by a s i n g l e hyaena, whereas an average of 11.5 clan members hunt zebra (Kruuk 1972). In wolves, caribou are usually k i l l e d by s i n g l e wolves or a p a i r ( C r i s l e r 1956, Haber 1977), however, larger prey such as moose are s u c c e s s f u l l y hunted only i n large packs (Mech 1966, Haber 1977). In Jasper, coyotes generally foraged for mice and ground s q u i r r e l s alone. Of the 80 times I observed coyotes foraging f o r small mammals, 80 percent involved a s i n g l e animal. In 193 contrast, two to four coyotes were involved i n 11 of 13 instances of attempted or successful ungulate predation. Thus, coyotes also appear to adjust group s i z e to the s i z e of prey hunted. Within the Hyaenidae we f i n d further evidence that group"foraging allows larger prey to be captured. There are three species i n t h i s family; the spotted hyaena, "the brown hyaena, Hyaena brunnea, and the s t r i p e d hyaena, Hyaena hyaena. A l l three species are approximately the same s i z e (Dorst and Dandelot 1970) and yet only the group-living spotted hyaena r e g u l a r l y preys on large ungulates (Kruuk 1972, 1975). The other two species are s o l i t a r y and feed mainly on small prey (Kruuk 1975, 1976, Eaton 1976, M i l l s 1978). As noted above another advantage of grouping may be increased a b i l i t y to defend the feeding area or concentrated food. In coyotes, t h i s applied p a r t i c u l a r l y to the use of ungulate carcasses. Packs (>2 animals) made s i g n i f i c a n t l y more e f f i c i e n t use of carcasses than s i n g l e residents or v i s i t o r s . This was evident both i n access to food and i n time spent feeding. This argument includes both i n t r a - and i n t e r - s p e c i f i c competition for l i m i t i n g resources. Since most carnivores r e a d i l y scavenge and competition f o r ungulate carcasses may be intense (Kruuk 1972, Sc h a l l e r 1972, t h i s study), the a b i l i t y to defend food against conspecifics or other species may favour grouping. This i s true regardless of whether or not prey i s captured. I f , for example, a carnivore were the largest member of a predator g u i l d , but smaller subordinate species grouped,, the l a r g e r species might be prevented from defending i t s own k i l l s or s t e a l i n g from others. This i n turn might favour grouping i n the larger species. There i s some support 194 for t h i s hypothesis. Mech (1970) notes that a s i n g l e g r i z z l y bear can exclude a wolf from i t s k i l l , but even a small pack of wolves w i l l success-f u l l y defend a carcass against t h i s species. Schaller (1972) found that one l i o n could not defend a k i l l against a pack of hyaenas, but two or more could. However, equally important was the f i n d i n g that grouped l i o n s s u c c e s s f u l l y s t o l e food from packs of hyaenas, but one l i o n could not. Food stolen from hyaenas forms the majority of the l i o n ' s d i e t i n some areas (Kruuk 1972). It has also been suggested that group foraging i n b i r d s reduces the overlap i n foraging areas allowing prey d e n s i t i e s to recover before the area i s searched again (Short 1961, Morse 1971). F i n a l l y , Hechrotte (1967) and Crook (1970) suggest that communal foraging increases the area searched for food. The extent to which these mechanisms may have favoured group foraging i n coyotes i s unknown. Schoener (1971), Alexander (1974) and others have argued that group foraging per se may be of neutral s e l e c t i v e value i n s i t u a t i o n s where animals aggregate i n proportion to the concentration of food. I suggest that coyote aggregations at ungulate carcasses (Murie 1940, Camenzind 1978, t h i s study) represent t h i s type of foraging group. Camenzind reported aggregations of up to 22 coyotes, including transients, resident p a i r s and packs, near elk c a r r i o n i n winter. As i n Jasper, where up to 14 coyotes were observed i n winter aggregations, such groups displayed no s o c i a l organization but were characterized by frequent agonistic behaviour as animals competed for p o s i t i o n at the food. In both areas, aggregations broke up into singles or pairs when coyotes foraged 195 for more dispersed food. Similar aggregations near c a r r i o n are commonly reported i n j a c k a l s , Canis aureus and C_. mesomelas (Estes 1967, van Lawick and van Lawick-Goodall 1970). Although common at ungulate carcasses i n winter, these aggregations are ephemeral with group s i z e and membership changing r a p i d l y and there i s no doubt that they are q u a l i t a t i v e l y d i f f e r e n t than the coyote packs observed. However, as"suggested by Kleiman and Eisenberg (1973), i t i s possible that aggregations may have been "an important step potentiating the development of cooperative hunting." This would be p a r t i c u l a r l y true i f re l a t e d animals cooperated i n the defense of concentrated food (see below). I conclude that group foraging increases the feeding e f f i c i e n c y of coyotes by allowing them access to large mammal prey. Coyote packs may be able to capture mule deer more e f f e c t i v e l y than s i n g l e animals or p a i r s . But, i n t h i s study, cooperative defense of ungulate carcasses from co n s p e c i f i c competitors i s l i k e l y the major benefit derived from group foraging. 196 10 SOCIAL ORGANIZATION IN RELATION TO ECOLOGY Recent research i n behavioural ecology has emphasized the influence of a species ecology on the evolution of s o c i a l behaviour, population disper-sion, and s o c i a l structure (see Crook 1970a for review). Thus, many behavioural c h a r a c t e r i s t i c s are viewed as adaptations to important s e l e c t i o n pressures in the environment (Crook 1964). For example, Crook (1964) found that differences i n s o c i a l organization among Ploceine weaver birds were correlated with contrasts i n food supply and feeding methods. Dispersed breeding, a c t i v e pursuit i n courtship and monogamy were.'/-.correlated with a r e l a t i v e l y abundant and persistent insect food that occurred p r i m a r i l y i n f o r e s t s . Savannah species, i n contrast, displayed f l o c k i n g , r e l a t i v e l y s t a t i c courtship i n colonies and polygamy; t h i s being an adaptation to a seasonally f l u c t u a t i n g food resource. A number of other studies have also demonstrated a r e l a t i o n s h i p between a s p e c i e s ' s o c i a l organization and i t s ecology. These include the following diverse taxonomic groups: I c t e r i d s (Orians 1961), hyaenas (Kruuk 1975, 1976, M i l l s 1978), antelope (Jarman 1974), marmots (Barash 1974, Heard 1977), canids (Kleiman 1972, Kleiman and Eisenberg 1973), primates (Crook 1970b, Eisenberg, Muckenhirn, and Rudran 1972), and kangaroos (Kaufmann 1974). In each of these studies, e x p l o i t a t i o n of food and/or predation were regarded as the primary s e l e c t i o n pressures which have shaped s o c i a l organization. The socio-ecology (Crook 1970a) of coyotes has received l i t t l e a t tention. Coyotes have been studied i n many d i f f e r e n t environments, but few authors have rel a t e d s o c i a l organization to contrasts i n ecology. 197 Consequently, few r e l i a b l e comparative data are a v a i l a b l e and attempts at a synthesis are premature. However, I believe enough i s known to begin to r e l a t e coyote s o c i a l structure to environmental differences. The evolutionary basis of canid s o c i a l organization i s the bond between a heterosexual adult p a i r , which r e s u l t s i n a monogamous mating system and considerably parental investment by both the male and female (Kleiman 1977, Kleiman and Brady 1978). The p a i r bond may be l i m i t e d to or strongest during breeding and caring of young as i n the maned.wolf, Chrysocyon brachyurus (Kleiman 1972) and the red fox (Kleiman 1967, Fox 1975), or i t may be permanent as i n the coyote (Kleiman 1977, t h i s study) and other canids (van der Merwe 1953, Egoscue 1962, K i l g o r e 1969, Haber 1977). Kleiman and Brady (1978) suggested that i n species with a strong p a i r bond, such as the coyote, v a r i a t i o n i n group s i z e and s o c i a l structure r e s u l t s from differences i n the age of d i s p e r s a l of young. In coyotes, young usually over-winter with t h e i r parents but disperse at the onset of the next breeding season (Fox 1975). Bekoff (pers. comm.) observed that coyotes i n Rocky Mountain National Park, Colorado, were s o l i t a r y ; about 95 percent of sightings were of si n g l e animals. In t h i s population subadults dispersed i n t h e i r f i r s t winter. On the Turnbull National W i l d l i f e Refuge, Washington, K e l l y and E l t o n (unpubl. manuscript) reported only the adult p a i r and pups at four coyote dens suggesting heavy m o r t a l i t y but more l i k e l y complete d i s p e r s a l of the previous year's l i t t e r . In some populations, not a l l coyotes disperse during t h e i r f i r s t year. In Jasper, although some coyotes dispersed at age 7-11 months, others stayed with t h e i r parents for two or more years. Ryden (1974) and Camenzind 198 (1978), both working i n Wyoming, found that young coyotes commonly delayed d i s p e r s a l and a s s i s t e d i n rearing t h e i r parents' o f f s p r i n g . Yearlings have also been observed at t h e i r parents' den i n Yellowstone National Park (Robinson and Cummings 1947) . Jackals and coyotes are s i m i l a r i n s i z e and ecology. Thus, i t i s not s u r p r i s i n g that i n golden j a c k a l s , young have been observed with t h e i r parents at the den following the b i r t h of a subsequent l i t t e r (van Lawick and van Lawick-Goodall 1970). Also, Moehlman (1976, unpubl-; i n Kleiman. and Brady 1978) observed that nonbreeding black-backed j a c k a l s , thought to be young from a previous l i t t e r or s i b l i n g s of the breeding p a i r , commonly as s i s t e d i n rearing the young. Delayed d i s p e r s a l of young i s most pronounced i n the wolf,and Cape hunting dog (Lycaon p i c t u s ) ; each a highly s o c i a l species. In these species, the young are incorporated into an extended family and d i s p e r s a l , when i t occurs, does not take place u n t i l wolves are age two or three (Frame and Frame 1976, Zimen 1976). V a r i a t i o n i n d i s p e r s a l appears to account f o r differences i n s o c i a l structure of coyote populations. But, why should some young disperse and others not? Bekoff (1977b) has addressed t h i s question i n terms of the behavioural ontogeny of l i t t e r mates (see Discussion, Chapter 7). Here I consider e c o l o g i c a l factors which might a f f e c t d i s p e r s a l of young. F i r s t , l e t us review the consequences of d i s p e r s a l on the f i t n e s s of i n d i v i d u a l young. Those i n d i v i d u a l s who disperse as soon as they are p h y s i o l o g i c a l l y able to reproduce increase the p r o b a b i l i t y of maximizing t h e i r reproductive output than do l a t e dispersers. Since coyotes generally mature at 10-11 199 months, they should normally disperse i n t h e i r f i r s t winter. However, higher m o r t a l i t y of e a r l y dispersers may be a strong opposing s e l e c t i o n pressure. But, o f f s p r i n g that delay d i s p e r s a l face competition for resources i n c l u d i n g mates from t h e i r parent and s i b l i n g s . Parents, of course, also have an i n t e r e s t i n the d i s p e r s a l of t h e i r young f o r the i n c l u s i v e f i t n e s s (Hamilton 1964) of parents i s based on the s u r v i v a l of current and previous o f f s p r i n g . By allowing t h e i r o f f s p r i n g to a s s i s t i n r e a r i n g subsequent young, parents may increase t h e i r f i t n e s s i n two ways. F i r s t , parents with helpers o f t e n r a i s e more young than those without. This has been documented i n group breeding b i r d s (Rowley 19.65,^ 'Wo-If-enden 1975) and r e c e n t l y i n black-backed j a c k a l s (Moehlman 1978). Second, the chance of s u r v i v a l of those young that delay d i s p e r s a l may be increased through the experience gained by extended a s s o c i a t i o n with t h e i r parents. Consequently, the p a t t e r n of d i s p e r s a l of young, and thus the s o c i a l organ-i z a t i o n of a population, w i l l be determined by e c o l o g i c a l f a c t o r s which a f f e c t the b e n e f i t / c o s t r a t i o to parents and young of having young stay with t h e i r parents. Young should disperse only when the cost to an i n d i v i d u a l and i t s parents exceed the b e n e f i t s of group l i v i n g . In Jasper, g r o u p - l i v i n g and t e r r i t o r i a l i t y were associated with a preponderance of ungulate food i n the coyote d i e t , e s p e c i a l l y i n winter (67 percent). S i g n i f i c a n t l y , these same t r a i t s were found i n coyotes i n Jackson Hole, Wyoming (Weaver 1977, Camenzind 1978)- I n t h i s population, e l k c a r r i o n formed the s i n g l e most important winter food (74 percent) of packs and deer and other ungulates accounted f o r another 7 per-cent of the d i e t . I i n d i c a t e d i n Chapter 9 that the p o s s i b l e advantages 200 coyotes derived from group l i v i n g were increased predation success on mule deer and more e f f i c i e n t defense and u t i l i z a t i o n of an ungulate carcass against i n t r a s p e c i f i c competitors. In the Wyoming population, ungulate food was mostly scavenged; therefore group-living coyotes probably benefited from cooperative defense of food, but t h i s was not mentioned by Camenzind. There are s t r i k i n g s i m i l a r i t i e s i n the d i e t and s o c i a l organization of group-living coyotes, wolves and Cape hunting dogs; best known as predators of large mammals. For example, a l l three species combine s o c i a l i t y with the use of large mammals as a major food. In the l a t t e r two species, t h i s may be necessary because of t h e i r large s i z e . The wolf, weighing from 27 to 54 kg (Mech 1970), consumes an estimated 2.6 kg (Mech and Frenzel 1971) to 6.3 kg (Mech 1966) of food per wolf per day. For hunting dogs, weighing 18 to 27 kg, estimated rates of consumption are 2.0 to 4.0 kg per dog per day (Estes and Goddard 1967). Thus, i t seems u n l i k e l y that e i t h e r species could e f f i c i e n t l y meet t h e i r energy requirements feeding on small rodents and lagomorphs. On the other hand, large ungulate prey are not e a s i l y captured by one animal. Coyotes, however, can e a s i l y l i v e on small prey. Why, then, do coyotes group to e x p l o i t large prey i f costs are incurred i n group-living? Given a choice between foods containing equal amounts of energy and assuming equal cost of feeding, coyotes would meet t h e i r energy requirements more quickly by feeding on large foods than on small ones. More time would then be a v a i l a b l e f or other a c t i v i t i e s such as t e r r i t o r i a l behaviour, which might increase the food a v a i l a b l e i n the t e r r i t o r y by l i m i t i n g s t e a l i n g by neighbours. Or, coyotes might rest more of the time, thereby reducing energy expenditures. Since a l l animals must acquire energy to meet t h e i r mainten-ance requirements before they may grow or reproduce, the f i t n e s s of coyotes 201 with greater net energy gain may be increased. Schoener (.1971) reviewed numerous examples which indi c a t e a causal r e l a t i o n between food intake and reproductive output. I t follows, then, that given a high density of ungulates, group-foraging coyotes should have a greater net energy gain per unit of feeding than s o l i t a r y animals. This would be p a r t i c u l a r l y true i f there i s considerable competition f o r ungulate food i n winter suggesting that hunting small mammals involves greater cost than the b e n e f i t derived from competing for ungulate food. Evidence of food competition among coyotes i n Jasper includes 1) i n d i v i d u a l s being prevented from feeding on ungulate carcasses through aggressive behaviour of residents or other v i s i t o r s , and 2) coyotes often waiting unsuccessfully for hours to gain access to carcasses monopolized by resident packs. However, I have no evidence that the s u r v i v a l of s o l i t a r y coyotes was l e s s as a consequence of t h i s competition. One needs to be cautious i n making arguments about the f i t n e s s of i n d i v i d u a l s . I t i s important to r e a l i z e that many factors can a f f e c t an i n d i v i d u a l ' s i n c l u s i v e f i t n e s s . In studies of adaptation, i f a s t r u c t u r a l or behavioural t r a i t can be shown or inferred- to increase i n c l u s i v e genetic f i t n e s s then t h i s t r a i t i s said to have adaptive value or adaptive s i g n i f i -cance. Although, i n c l u s i v e f i t n e s s has emerged as "the ultimate currency i n which to evaluate a l t e r n a t i v e behaviours" (Barash 1975), i n p r a c t i c e i t i s seldom possible to measure f i t n e s s . We must often s e t t l e f o r demonstrat-ing that a behaviour helps the animal to increase rate of net energy gain or avoid, predation. We, then, i n f e r (as above) that these behaviours w i l l probably increase f i t n e s s . 202 To test the hypothesis that group l i v i n g allows coyotes to capture larger prey or use i t more e f f i c i e n t l y once a carcass i s found, one could 1) experimentally manipulate the s i z e of prey a v a i l a b l e to coyotes within a population or 2) compare s o c i a l structure between populations where the average s i z e of prey eaten by coyotes d i f f e r s markedly. Based on the r e s u l t s i n Jasper, (this study) and i n Wyoming (Camenzind 1978), I suggest that coyotes l i v i n g p r i m a r i l y on small prey disperse during t h e i r f i r s t winter and that the nuclear family i s the basic unit of s o c i a l organization. To my knowledge only one study has been conducted on a population of t h i s type. Bekoff (pers. comm.) observed that s o l i t a r y coyotes i n Rocky Mountain National Park, Colorado, fed mainly on small rodents. Bekoff believed that young dispersed during t h e i r f i r s t winter and that the nuclear family was the basis of s o c i a l organization i n the population. Thus there i s some support for the hypothesis. As i n any natural experiment, differences between these populations other than s i z e of prey eaten may have affected coyote s o c i a l organization. A comparison between- populations, i s v a l i d only i f the abundance of a v a i l a b l e food was equal i n both populations. It i s c l e a r , f o r example, that pride s i z e and cohesiveness i n l i o n s i s very dependent on food supply. In the Serengeti, an area of high ungulate density, Schaller (1972) found that the average number of l i o n s i n a pride.was 15 (n = 14, range of 4 to 37). However, i n the dunes of the K a l i h a r i region where ungulate biomass i s low, l i o n s occur i n small prides ranging from 2 to 13 animals (x = 5.2, n = 6) or are s o l i t a r y ( E l o f f 1973). Population density might'also a f f e c t the l e v e l of coyote s o c i a l i t y , independent of the s i z e of prey eaten. The e f f e c t of population density on 203 the s o c i a l structure of rodents has been reviewed by Archer (1970). Of p a r t i c u l a r i n t e r e s t i s the f i n d i n g that i n Mus muscuius, s o c i a l i t y increases with increasing population density. Anderson (1961) concluded that i n p a r t i c u l a r l y favourable environments, the t e r r i t o r i a l behaviour of house mice breaks down and an extensive sharing of space occurs. In wolves, pack s i z e increases to an asymptote with population density (Rausch 1967, Mech pers. comm. i n Zimen 1976). Population density has also been proposed as a fac t o r a f f e c t i n g i n t r a s p e c i f i c v a r i a t i o n i n the s o c i a l structure of c e r t a i n Old World monkeys, Cercopithecidae (see Crook 1970b f or review). Another p o s s i b i l i t y i s that coyote s o c i a l i t y i s affected by the spatio-temporal d i s t r i b u t i o n of food. A food resource that i s clumped and unpredictable i n time and space, may be located too infrequently by a si n g l e predator to be u t i l i z e d ; however, the c o l l e c t e d searching e f f o r t s of a group may allow i t s e x p l o i t a t i o n ; For example, chimpanzee and spider monkey groups may s p l i t and reassemble at abundant food to better locate i r r e g u l a r l y seasonal f r u i t (Reynolds and Reynolds 1965, Crook 1970b) and various birds use f l o c k i n g to e x p l o i t temporary food (Rand 1954, Crook 1964, Kahl 1964, Ward 1965, Horn 1968, Krebs 1974, Davies 1976). Human e x p l o i t a t i o n may also influence the s o c i a l structure of c a r n i -vore populations. Kleiman and Brady (1978) suggest that f i e l d studies of population and behavioural ecology w i l l be severely Biased by human i n t e r -ference and persecution of red foxes, coyotes and wolves.. Lack (1965) emphasized the importance of studying evolutionary ecology i n "the natural habitat of a species, as many e c o l o g i c a l adaptations may be obscured i n 204 man disturbed environments. Storm et a l . (1976) found that the majority of a l l red fox deaths i n Iowa each year are due to human a c t i v i t i e s . The e f f e c t of human e x p l o i t a t i o n on the population biology of coyotes (Knowlton 1972) and wolves (Rausch 1967, Mech 1970) are well documented. In unexploited wolf populations only 60 percent of adult females breed annually (Pimlott et a l . 1969), whereas Rausch (1967) found.that 89 percent of females were pregnant where wolves were heavily exploited. Rausch (1967) reported that average pack s i z e i n wolves was markedly reduced under the influence of man. I t i s l i k e l y that our i n a b i l i t y to c l e a r l y characterize the s o c i a l structure of coyotes i s due i n part to heavy human e x p l o i t a t i o n . There are numerous accounts of coyote packs by r e l i a b l e early n a t u r a l i s t s and explorers (Dobie 1949). Thomas Say, who f i r s t c l a s s i f i e d the coyote as Canis latrans i n 1823, wrote " P r a i r i e wolves are by f a r the most numerous of our wolves, and often unite i n packs f o r the purpose of chasing deer." (quoted i n Dobie, 1949). The f a i l u r e to recognize the r e l a t i o n s h i p between prey s i z e and coyote s o c i a l structure despite the observed v a r i a t i o n i n s o c i a l structure and s i z e of prey eaten (Table XXIX), i s probably due to human e x p l o i t a t i o n of.these populations. In Table XXIX, I have calculated the mean percent frequency of each food class i n the d i e t for Jasper and a l l other areas combined. The 22 data sets from 18 studies are derived from stomach-content and f e c a l a n a l y s i s . In most of these studies, young ungulates (class 3 prey) were lumped with adults. Thus, the percentage of class 4 prey i n summer d i e t s i s an overestimate i n these cases. Class 4 prey occur i n approximately 22 percent of the die t i n the combined sample i n summer and about 38 percent i n winter. In contrast, the die t i n Jasper consists of 30 percent class 4 prey i n summer and 61 percent i n winter. Table XXIX. Percent frequency of food classes 1 to 4 (see Table 19) i n coyote diets based on 18 studies. Food class 1 2 3 4 Source Summer Winter Summer Winter Summer Winter Summer Winter Bond (1939) a' b 42 14 23 20. Brown (1977) b 69 36 5 10 13 15 13 35 F e r r e l e t a l . (1953) , c 25 32 26 10 19 15 31 39 18 18 9 6 28 32 42 36 23 27 13 28 23 35 42 16 16 10 4 53 43 18 39 15 27 18 13 44 33 21 24 Fich t e r et a l . (1955) b 55 43 25 32 5 9 15 16 Gier (1968)a7b 29 5 44 . 22 Gipson (1972) b 37 39 11 2 41 28 11 28 Knowlton (1964) b 28 29 16 8 15 ' 19 37 41 Korschgen (1957) b 7 15 29 19 42 53 22 13 Mathwig (1973) b 11 0 71 21 14 Meinzer et a l . (1975) 62 4 11 Murie (1940) 46 27 9 15 N e l l i s and Keith (1976) 23 5 11 61 Ozoga and Harger (1966) 19 19 11 51 Richens and Hugie (1974) b 15 0 26 42 Weaver (1977) b 39 !& 48 3 4 1 9 81 X 34.0 24.4 17.6 9.4 23.5 26.0 22.1 37.5 S.E. 4.74 2.54 3.06 2.10 4.17 4.78 2.55 4.67 This study , 26 23 15 2 22 5 32 67 Cowan (1943) b 15 11 30 6 12 13 43 70 Hatler (1945) 23 21 22 2 40 30 14 45 X 21.3 18.3 22.3 3.3 24.7 16.0 29.7 60.7 S.E. 3.28 3.71 4.33 1.33 7.59 7.37 8.46 7.89 Both summer and winter samples included. Ungulate young not l i s t e d separately i n summer. Five d i f f e r e n t regions of C a l i f o r n i a represented. 20.6 Although, average prey s i z e eaten i n Jasper i s greater than f o r most other populations, i t i s clear that there i s considerable v a r i a t i o n i n the proportion of class 4 foods (ungulates) i n the d i e t . Thus, given the average s i z e of a coyote pack (4 to 5 animals) and the high l e v e l of man-related mortality i n these populations, i t i s not s u r p r i s i n g that coyote packs are not reported, even where ungulates accounted for 40 to 60 percent of the d i e t . I t would be an Interesting"test of the hypothesis to h a l t the hunting of populations where ungulates comprise a major part of the d i e t . Would coyote group s i z e increase? Another factor which may favour s o c i a l i t y i n coyotes i s reduced r i s k of predation e i t h e r as adults or young. A number of carnivores are known to prey on young and adult coyotes and jackals (Cowan 1947, Dobie 1949, Young and Jackson 1951, van der Merwe 1953, van Lawick and van Lawick-Goodall 1970, Camenzind 1978). Unfortunately, most of t h i s information i s anecdotal and thus the importance of predation as a f a c t o r i n f l u e n c i n g grouping i n coyotes cannof.be assessed at t h i s time. Wilson (1975) discounted r i s k s from predation as important i n the evolution of group l i v i n g i n larger carnivores. However, Eaton (1976) points out that a l l the larger f e l i d s studied to date are k i l l e d as adults by competitors and t h e i r young incur predation. Eaton (1976) suggests that, although the female l i o n separates from the pride at p a r t u r i t i o n , the l i t t e r i s normally introduced to the group as soon as the young are able to t r a v e l with the pride and benefit from group defense. There i s some support for t h i s idea. In Nairobi Park, the major predators of young l i o n s are v i r t u a l l y absent and female l i o n s group more rloosely than elsewhere (Rudnai 1973). 207 In small and medium s i z e c a r n i v o r e s group l i v i n g to reduce predation may be more common. For example, the banded mongoose, Mungos nuingo, i s a sma l l v i v e r r i d that l i v e s i n troops of up to 32 i n d i v i d u a l s . I t forages s i n g l y f o r i n s e c t s and small v e r t e b r a t e s . In response to predators, members of a troop, j o i n i n a t i g h t bunch w i t h mouths p o i n t i n g i n a l l d i r e c t i o n s , g i v i n g the appearance of one l a r g e organism defending i t s e l f (Rood 1973, Neal 1970). Grouping i n k i t foxes (Egoscue 1962) might a l s o be i n t e r p r e t e d as a s t r a t e g y to reduce pre d a t i o n . F o r t u n a t e l y , many of the problems discussed do not apply to t h i s study. The Jasper p o p u l a t i o n has not been s e r i o u s l y hunted s i n c e the 1950s, but continued road deaths probably decrease pack s i z e below that of t r u l y p r i s t i n e c o n d i t i o n s . Coyote d e n s i t y estimates (Chapter 5) i n Jasper agree w i t h average d e n s i t i e s over much of t h i s species range. While c e r t a i n prey, such as Columbian ground s q u i r r e l s , are clumped, they are p r e d i c t a b l e i n time and space as are the ungulate species. I t i s , t h e r e f o r e , u n l i k e l y that coyotes l i v e i n packs f o r t h i s reason. P r e d a t i o n of coyotes by wolves and mountain l i o n s was r a r e and bear pre d a t i o n was not found. Thus we are l e f t w i t h group f o r a g i n g as an adaptation to l a r g e prey s i z e as the hypothesis which best e x p l a i n s the observations i n Jasper ( t h i s s t u d y ) , and Wyoming (Camenzind 1978). Increased f o r a g i n g e f f i c i e n c y i s almost u n i v e r s a l l y accepted as a d r i v i n g f o r c e favouring s o c i a l i t y i n the l a r g e carnivores (Kleiman"and.Eisen-b'.ergV- 1973, Alexander 1974, Kruuk 1975, Wilson, 1975). This has lead Fox (1975) to propose three types of canid s o c i a l s t r u c t u r e s based l a r g e l y on the extent to which group f o r a g i n g i s observed and on~the s t r e n g t h of 208 r e l a t i o n s h i p s between adults and between adults and t h e i r o f f s p r i n g . The red fox i s used as an example of the Type I or s o l i t a r y s o c i a l structure. In t h i s type, adult males and females form a temporary p a i r bond during the period of breeding and rearing young; the parent-young bond i s weak by f a l l when young disperse from t h e i r n a t a l range. Correlated with t h i s s o c i a l structure i s a s o l i t a r y mode of hunting i n which small rodents, f r u i t s and lagomorphs are the p r i n c i p a l prey (see Abies 1975, Lloyd 1975 for reviews). In Type II canids (e.g. coyotes), the adult p a i r bond and parent-young bond p e r s i s t outside of the breeding-rearing season. Offspring stay with t h e i r parents through t h e i r f i r s t winter and sometimes longer. Type II canids are e f f i c i e n t s o l i t a r y hunters of small prey, but are capable of hunting i n pair s or as a family u n i t . Fox uses the wolf as an example of a Type III"or pack s o c i a l structure i n which large ungulates are captured members of an extended family. Kleiman and Brady (1978) argue that although the c o r r e l a t i o n between hunting strategy and s o c i a l structure may be us e f u l i n understanding the evolution of group s i z e i n the larger canids, i t i s misleading when broadly applied to other canid species. For example, they note that k i t foxes (Egoscue 1962), swift foxes, Vulpes velox (Kilgore 1969), bat-eared foxes, Octocyon megalotis (Hendrichs 1972, c i t e d i n Kleiman and Brady 1978), and crab-eating foxes, probably l i v e i n stable pairs or small family groups throughout the year, and yet these species hunt s i n g l y and l i v e mainly on small prey. Fox's c l a s s i f i c a t i o n f a l l s short ult i m a t e l y because of i t s f a i l u r e to recognize the mu l t i v a r i a t e nature of in t e r a c t i o n s between a species and i t s environment. Future attempts w i l l have to consider the r o l e of (1) i n t r a - and i n t e r - s p e c i f i c food competition, (2) predation, 209 (3) other food dimensions such as absolute abundance and spatio-temporal d i s t r i b u t i o n , and (4) population density i n shaping canid s o c i a l organization. There i s also evidence that any theory of s o c i a l organization based s o l e l y on the concept of adaptation to contrasts i n ecology may f a l l short because of random events i n the evolutionary h i s t o r y of a s i n g l e or group of species. A small evolutionary change i n the behaviour of i n d i v i d u a l s may be amplified into a major e f f e c t on s o c i a l structure (Wilson 1975). As an example, consider the d i f f e r i n g s o c i a l organizations of the c l o s e l y r e l a t e d o l i v e baboon, Papio ariubis, and the hamadryas baboon, P_. hamadryas. The hamadryas male guards females throughout the year, whereas the o l i v e male appropriates females only around the time of t h e i r estrus. Although s l i g h t , t h i s d i f f e r e n c e alone i s enough to account f o r profound changes i n s o c i a l structure (JCummer / 1971) . It i s now recognized that, at l e a s t as frequently as morphological characters or s i g n a l l i n g behaviour, s o c i a l organization may show considerable v a r i a t i o n over a species range. For example, Kruuk (1972) demonstrated a change i n s o c i a l organization of spotted hyaenas from a t e r r i t o r i a l group-hunting type i n the Ngorogoro Crater to s o l i t a r y hunting, generally without t e r r i t o r i e s , on the adjacent Serengeti P l a i n s . He r e l a t e d these differences to the hyaena's food. In Ngorogoro Crater, the hyaena's preferred prey are permanent residents and reach high d e n s i t i e s ; thus, t e r r i t o r i a l defense by members of a clan i s economical. On the other hand, the lower absolute density and migratory behaviour of the hyaena's food supply on the Serengeti Plains made attachment to and defense of an area unfeasible. Haber (1977) showed that neighbouring wolf packs may d i f f e r markedly i n s o c i a l structure. Evidence from Haber (1977) and Mufie (1944) 210 indicated that more than one l i t t e r per year was common i n the Toklat Pack between the 1930s and 1970, whereas one l i t t e r only was t y p i c a l i n the Savage Pack. Haber (1977) a t t r i b u t e d t h i s to differences i n food leading to differences i n the type of pack s p l i t t i n g . The Toklat Pack u t i l i z e d seasonally abundant caribou :and low density resident moose and sheep populations. This led to the pack s p l i t t i n g into several groups i n winter and each containing experienced mature, wolves. In contrast, the Savage Pack's prey were permanent residents that reach,high d e n s i t i e s . Pack s p l i t t i n g i n t h i s pack was characterized by immature wolves only separating from the mature adults. Although more work i s needed, i t seems clear that coyote s o c i a l structure i s also affected by the nature of i t s food. In p a r t i c u l a r , d i f f e r -ences i n average s i z e of prey eaten account f o r much of the v a r i a t i o n i n s o c i a l c s t r u c t u r e . Group l i v i n g i s correlated with high ungulate density i n both Jasper ( t h i s study) and Wyoming (Camenzind 1978). Coyote packs cons i s t i n g of r e l a t e d adults, yearlings and young fed p r i m a r i l y on mule deer and elk. Although much of t h i s food was scavenged, mule deer predation may have contributed s i g n i f i c a n t l y to the d i e t of coyotes i n Jasper. In contrast, coyotes are s o l i t a r y or l i v e i n stable p a i r s where mostly small rodents and lagomorphs are eaten. V a r i a t i o n i n coyote s o c i a l structure has led to a maximizing of i t s resource base. S o c i a l i t y i s a strategy allowing access to large prey without s a c r i f i c i n g the advantage that smaller body s i z e gave i t i n the e f f i c i e n t use of small prey. 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S o c i a l dynamics of the wolf pack, p. 336 - 362. In M. W. Fox (ed.), The Wild Canids: Their Systematics, Behavioral Ecology and Evolution. Van Nostrand Reinhold, New York. v . . . 1976. On the regulation of pack s i z e i n wolves. Z. Tierpsychol. 40: 300-341. 225 Appendix I Aspects of age determination i n coyotes Two methods are commonly used to age coyotes: tooth-wear patterns of i n c i s o r s and canines (Gier 1968, unpubl. data) and, incremental cementum annuli (Linhart and Knowlton 1967). The f i r s t method simply requires a v i s u a l inspection of the teeth. Thus, i t allows rapid age determination of l i v e animals i n the f i e l d . The p r i n c i p a l disadvantage of t h i s method i s that there i s l i k e l y geographic v a r i a t i o n i n the rate of tooth wear depending on the types of food eaten. In contrast, to use cementum annuli for age determination, teeth are h i s t o l o g i c a l l y prepared f o r microscopic examination. Although t h i s i s expensive and time consuming i t i s generally regarded as the most r e l i a b l e method. A tooth must be extracted to use t h i s method. This presents a problem p a r t i c u l a r l y since the canine i s preferred and i t i s l i k e l y that removing a canine from an animal that i s to be released would have harmful consequences. There are several a l t e r n a t i v e solutions to t h i s problem. One i s to extract the f i r s t premolar, a small and l a r g e l y non-functional tooth, and proceed with h i s t o l o g i c a l preparation f o r cememtum annuli a n a l y s i s . Another s o l u t i o n i s to use the tooth wear method i n the f i e l d on l i v e coyotes, but r e c a l i b r a t e the index using the more accurate cementum annuli procedure on coyote teeth c o l l e c t e d from carcasses i n the same area. I used the second s o l u t i o n . The teeth of 30 coyotes c o l l e c t e d i n Jasper National Park were a v a i l a b l e 226 for study. The lower jaw of each animal was b o i l e d i n water for approximate-l y 1 hr to f a c i l i t a t e tooth extraction. Teeth were d e c a l c i f i e d i n a s o l u t i o n of 5 percent formic acid and 5 percent formaldehyde, time depending on age of teeth and then stored i n 70 percent alcohol. Tooth roots were sectioned l o n g i t u d i n a l l y at 24 y by a cryostat. Sections were^stained with Paragon s t a i n f o r 1-2 min, rinsed i n water and mounted on g e l a t i n i z e d s l i d e s . Sections were then a i r dried and mounted i n Permount. Some sections were stained- with D e l a f i e l d ' s hematoxylin stain~£or 10 min, rinsed i n water, "blued" i n a l k a l i n e water and treated as above. Two questions must be answered before we can use incremental cementum annuli as a method of age determination; when does the f i r s t annulus form, and how much time elapses between the formation of two successive annuli? The second question has been answered with c e r t a i n t y . Once the f i r s t annulus i s deposited, another i s formed i n each succeeding year (Linhart and Knowlton 1967). Regarding the f i r s t question, Linhart and Knowlton (1967) speculated that the f i r s t annulus was deposited between 20 to 25 months of age. Recently, A l l e n and Kohn (1976) reported that f i v e 18-month-old coyotes from North Dakota had 1 cementum annulus. In Jasper, one 15-month-old male had 1 c l e a r l y v i s i b l e annulus near the exterior of the cementum. In a second 15-month-old male, the f i r s t annulus appeared to be forming but was imcomplete. One 23-month-old male also had 1 c l e a r l y formed l i n e , however, a female of same age did not yet have a f u l l y formed f i r s t annulus. In t h i s case, the annulus appeared to be developing at the exterior edge of the cementum layer. Such considerable v a r i a t i o n i n the 227 time of f i r s t annulus formation could r e s u l t i n inaccurate aging. C l e a r l y , more de t a i l e d studies are needed. Several authors have studied the agreement i n the number of cementum annuli found i n canines and f i r s t .premolars taken from the same i n d i v i d u a l . Roberts (1978) found that these teeth agreed i n only 4 of 8 coyotes, whereas Knudsen (1976) reported that 73 percent of 43 i n d i v i d u a l s were assigned the same age based on both teeth. A canine and premolar were a v a i l a b l e from 21 of 30 coyotes c o l l e c t e d i n Jasper. Of these 21 i n d i v i d u a l s , the same age was assigned to 17 or 81 percent based on both teeth. These studies support the view that where possible the"canine should be used i n age determination. However, they also suggest that there i s geographic v a r i a t i o n i n the agreement between these two teeth. In c e r t a i n areas, such as Jasper, the premolar w i l l provide a reasonably accurate estimation of age. Coyotes were assigned the same age using tooth wear and cementum annuli methods 53.3 percent of the time (n =30). Tooth wear patterns were f a i r l y r e l i a b l e f o r some age classes. Only two of 11 subadult and y e a r l i n g coyotes were i n c o r r e c t l y aged by tooth wear. However, 67 percent of s i x age-two coyotes and 73 percent of 11 age-three-plus coyotes were i n c o r r e c t l y aged. Of four age-two animals i n c o r r e c t l y aged, age was underestimated twice and overestimated twice. For older coyotes, tooth wear generally underestimated age and the error increased with the age of the coyote. Therefore, i n Jasper, tooth wear patterns r e l i a b l y aged subadults and yearl i n g s , but not older animals. Appendix II.. Age, weight and body measurements of 19 male and 20 female a d u l t a coyotes from Jasper National Park. Males Age (yr.mon) Weight (kg) Body length-(cm) T a i l length (cm) Hind foot length (cm) CIO 5,8 12.2 94.0 30.5 19.1 C12 2,0 14.3 105.4 33.0 19.8 C13 1,1 14.1 94.0 36.8 19.8 C14 ^ « . l 12.2 92.7 33.0 18.0 C16 '•' 2,0 12.7 91.4 33.0 19 v 8 C18 1,3 12.7 88.9 33.0 18.8 C19 2,1 13.8 92.7 34.3 19.3 C20 1,1 14.1 91.4 34.3 19.1 C23 8,0 13.9 94.6 34.6 19.8 C25 3,0 14.1 92.7 33.3 19.3 C50 4,3 11.8 88.9 33.0 16.8 .RK5 2,6 11.8 76.2 31.2 18.5 RK7 2,7 7.5 76.2 30.5 17.8 RK12 . 3,7 11.1 78.7 36.8 17.8 RK15 2,9 8.2 78.7 34.3 18.8 RK17 4,0 10.9 84.7 33.3 18.0 RK18 4,8 11.8 88.9 33.0 19.3 RK30 1,9 12.2 94.0 35.1 19.1 RK33 4,8 15.0 94.0 35.6 19.3 Females Age Weight Body length T a i l length Hind foot (yr, mon) (kg) (cm) (cm) length (cm) C2 5,7 12.2 83.8 30.5 18.5 C3 8,7 12.7 . 91.4 35. i 19.1 C4 2,0 12.7 91.4 29. 2 19.1 . C5 2,8 10.0 83.8 30. 5 18.5 C6 10,3 12.9 91.4 34. 3 13.3 C l l 6,0 . 12.9 91.4 33.0 18.5 C15 3,5 12.0 92.7 33.0 19.1 C24. 1,0 .9.8 91.4 35. 6 17.8 C26 . 2,0 13.6 93.4 35. ,6 19.1 C27 3,0 12.2 91.4 30. ,0 18.5 RK4 5,10 10.4 88.9 31. .2 19.1 RK13 2,0 11.8 83.8 • 31. ,2 18.5 RK16 5,0 12.9 86.4 32, .5 18.5 RK20 5,10 12.7 91.4 34, .3 19.8 RK22 1,8 8.6 83.8 33, .0 . 18.3 RK26 4,2 . 10.4 68.6 33, .0 18.5 RK31 1,8 9.1 81.3 29 , 2 17.8 RK35 8,7 12.9 91.4 33 .0 " 18.8 RK36 1,7 11.7 91.4 31 .2 18.5 RK39 3,1 11.5 88.9 33 .0 18.3 OO a>12 months of age. Appendix I I I . Percent frequency of prey species i n the di e t of d i f f e r e n t coyote groups i n winter, 1974-75 and 1975-76 combined (A), and i n summer, 1974 to 19/b combined (B). Prey types Athabasca Pack (n = 101) Rocky River Pack (n = 40) Miette Snaring Pack Pack (n = 12) (n = 38) Palisades Pack (n = 47) Cinquefoil Pack (n = 58) Talbot Lake Pair (n = 21) Maligne Pack (n = 319) Aloes aloes Cervus canadensis Odocoileus hemionus Ovis canadensis Lepus americanus Castor canadensis Ondatra sibethicus Microtus sp. Peromyscus manioulatus Clethrionomys gapperi Phenacomys intermedins Spevmophilus columbianus Neotoma cinerea Tamiasciurus hudsoniaus Birds Fish Vegetation U n i d e n t i f i e d food 22 42 5 14 4 11 38 18 17 67 42 13 16 3 5 3 27 15 7 20 3 2 2 2 17 19 16 5 2 41 5 29 29 10 29 21 47 3 13 2 1 K3 ro VO B Prey types Athabasca Pack (n = 183) Rocky River Pack (n = 120) Miette Pack (n = 61) Snaring Pack (n = 154) Palisades Pack (n = 120) Cinquefoil Pack (n = 38) Talbot Lake Pair (n = 40) Maligne Pack (n = 131) Kinross Whistlers Pair Pack (n = 46) (n = 79) Alces aloes Cervus canadensis (ad) (young) Odocoileus hemionus (ad) " _ (young) Ovis canadensis Lepus americanus Spermophilus columbianus Castor canadensis Ondatra zibethicus Mierotus sp. Peromyscus maniculatus Clethrionomys gapperi Phenacomys intermedins Marmota galigata Tamiasciurus hudsonicus Neotoma oinerea Ursus americanus U n i d e n t i f i e d mammal Birds F i s h Vegetation Insects U n i d e n t i f i e d food 4 11 1 10 25 6 5 14 3 Tr 26 6 11 23 2 7 7 3 2 2 2 1 3 Tr 1 66 2 1 14 6 15 3 2 1 7 T r a Tr 1 27 31 6 18 2 25 18 1 3 2 13 2 1 1 1 3 1 Tr 3 18 5 7 5 5 29 4 Tr 5 3 Tr 35 8 3 3 5 3 Tr 10 2 •> 12 1 Tr 13 2 2 1 1 1 5 5 2 1 12 16 4 28 25 4 2 2 11 9 6 18 4 28 32 NJ OJ O Percent frequency < 1 percent. 

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