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The behaviour of Vancouver Island marmots, Marmota vancouverensis Heard, Douglas C. 1977

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THE BEHAVIOUR OF V&NCOUVEB I S L A N D HABBOTS , MARMOTA VANCOUVBRENSIS E . S c . , U n i v e r s i t y o f W a t e r l o o , 1973 A T H E S I S SUBMITTED I N P A R T I A L P U L P I L L M E N T OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF S C I E N C E i n THE FACULTY OF GRADUATE STUDIES (The D e p a r t m e n t o f Z o o l o g y ) We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e r e g u i r e d s t a n d a r d THE U N I V E R S I T Y OF B R I T I S H COLUMBIA N o v e m b e r , 1977 by DOUGLAS C . HEARD D o u g l a s C . H e a r d , 1 9 7 7 In p r e s e n t i n g t h i s t h e s i s in p a r t i a l f u l f i l m e n t o f the r e q u i r e m e n t s f o r an advanced degree at the 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 that the L i b r a r y s h a l l make 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 and 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 may be g r a n t e d by the Head o f my Department or by h i s r e p r e s e n t a t i v e s . It i s u n d e r s t o o d tha 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 not be 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 . Department o f The U n i v e r s i t y o f B r i t i s h Co lumbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date ? ^ 2 ^ 0 7 ^ ABSTRACT I studied the s o c i a l behaviour of the Vancouver Island marmot, Marraota Vancouverensis, during the summers of 1973 and 197*». V i r t u a l l y nothing was known about the behaviour of this species at the outset of t h i s study, Barash {1973b, 1974a) suggested that the s o c i a l behaviour and s o c i a l organization of marmot species was determined by the severity of the environment (the vegetative growing season) and i t s e f f e c t on the growth rate of marmots. He predicted that marmot species l i v i n g i n short growing season environments would be highly s o c i a l but that s o c i a l tolerance would decrease as the growing season increased. The objective of th i s study was to test t h i s hypothesis by observing the s o c i a l behaviour of Vancouver Island marmots and comparing t h i s to the length of the vegetative growing season. J3« vancouyerensis i s endemic to Vancouver Island, B r i t i s h Columbia. The o r i g i n a l colonizers of t h i s species probably crossed to Vancouver Island via land connections that existed during the I l l i n o i a n g l a c i a l period, approximately 100,000 years ago, and survived subseguent g l a c i a l maxima on nunataks and coastal refugia or both. Vancouver Island marmots have been isolated from mainland forms for a length of time (10,000 to 100,000 years) s u f f i c i e n t to show s p e c i f i c evolutionary adaptations to t h e i r Vancouver Island environment. Vancouver Island marmots l i v e in small colonies in the i i i subalpine parkland. S o c i a l groups consisted of one adult male, one adult female, and variable numbers of two-year-olds, yearlings, and infants. S o c i a l groups were highly integrated with a large amount of communication occurring among colony members. Alarm c a l l s were given i n response to potential predators and could be heard over the whole colony. Short whistles were given i n response to a e r i a l predators (e.g. eagles) and long whistles were given i n response to t e r r e s t r i a l predators (e.g. black bears). Both c a l l s are narrow bandwidth sounds, a c h a r a c t e r i s t i c that makes them d i f f i c u l t to locate. The most common s o c i a l behaviour that occurred among colony members was a nose touching behaviour termed greeting. A l l age-sex classes of Vancouver Island marmots engaged i n greetings as well as other s o c i a l behaviour patterns i n about the same proportions. The vegetative growing season experienced by Vancouver Island marmots was approximately the same as that of M.,flaviventris but the s o c i a l behaviour of Vancouver Island marmots most c l o s e l y resembled M. oly_i£us, a species l i v i n g where the growing season i s much shorter. On t h i s basis I rejected Barash*s hypothesis that the length of the vegetative growing season i s s u f f i c i e n t to account for the v a r i a b i l i t y that Barash observed among marmot species. I suggest that vegetative growing season not be used as an index of growth rate but that the time taken to reach adult size be measured d i r e c t l y . The degree of s o c i a l tolerance i s p o s i t i v e l y correlated with the length of time reguired to reach maturity. i v TABLE OF CONTENTS A b s t r a c t . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . i i L i s t o f F i g u r e s v i L i s t o f T a b l e s v i i i A c knowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . x I n t r o d u c t i o n .. .. 1 S o c i o b i o l o g y and B a r a s h ' s H y p o t h e s i s . . . . . . . . . . . . . . . . . . . 1 V a n c o u v e r I s l a n d Marmots 4 Taxonomy and E v o l u t i o n a r y H i s t o r y . . . . . . . . . . . . . . . . . . . 5 Methods ... 10 S t u d y A r e a s 10 Methods o f O b s e r v a t i o n 10 A n a l y s i s o f S o c i a l B e h a v i o u r D a t a . . . . . . . . . . . . . . . . . . . . . . 14 T r a p p i n g and M a r k i n g 19 Measurement o f M i c r o c l i m a t e . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 V o c a l i z a t i o n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 R e s u l t s . . . . . . . . . . . . . . . . . . . . . . . . . . 22 H a b i t a t C h a r a c t e r i s t i c s 22 P h y s i c a l C h a r a c t e r i s t i c s 26 C o l o n y C o m p o s i t i o n 33 A c t i v i t y P a t t e r n s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 V o c a l i z a t i o n s .... ... ...... 44 W h i s t l e s . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Keeaws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... v .,... ..,. 54 R a p i d C h i r p s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Hisses .............................................. 57 T o o t h C h a t t e r s 57 Screams and G r o w l s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 S o c i a l Behaviour 59 Social Behaviour Patterns 59 Dominance Relationships 63 The Frequency of Social Behaviour Patterns .......... 66 Interaction Sequences ............................... 72 Rates of Social Behaviour ........................... 74 Dispersion, T e r r i t o r i a l i t y , and Scent Harking ....... 82 Discussion ................ *............................... 91 vocalizations .......................................... 91 i n t e r s p e c i f i c Comparisons of Marmot Vocalizations .. 91 Altruism and Marmot Alarm C a l l i n g ................... 94 The Evolution of Hhistle Structure .................. 97 The Evolution of Whistle Function ................... 99 In t e r s p e c i f i c Comparisons of Marmot Social Behaviour ..102 A Test of Barash's Hypothesis ....................... 102 Literature Cited .............. ................ ....,....... 115 Appendix I. A L i s t of Plant Species Found on the Haley Lake Study Area Indicating Those Species Known to be Eaten by Vancouver Island Marmots ....................... 125 Appendix I I . A L i s t of a l l Known Vancouver Island Marmot Specimens that have been Collected, Total = 30 ...... 129 v i LIST OF FIGURES Figure 1. Locations of Known Vancouver Island Marmot Colonies . 11 Figure 2. Photographs of the Haley Lake Study Area ....... 12 Figure 3. Photographs of Vancouver Island Marmot Habitat . 2 3 Figure 4. Seasonal Changes i n the Weights of Vancouver Island Marmots . . . . . , ;27 Figure 5. Seasonal Changes i n the Weights of Adult Marmots ..... . .. . . . ..... .. . ... . . ....... . . ..... .... 29 Figure 6. A c t i v i t y Budgets for May, a l l Animals Combined . 3 9 Figure 7. A c t i v i t y Budgets for June, a l l Animals Combined 40 Figure 8. A c t i v i t y Budgets for July, a l l Animals Combined 41 Figure 9. A c t i v i t y Budgets for August, a l l Animals Combined ........................... .................... 42 Figure 10. A c t i v i t y Budgets for September, a l l Animals Combined 43 Figure 11. Representative Sonogram of a Short Whistle ..... 45 Figure 12. Representative Sonogram of a Long Whistle ...... 46 Figure 13. A Comparison of the Length of Long and Short Whistling Sequences ...... .......... .................... 48 Figure 14. Representative Sonograms of Keeaws 55 Figure 15. Freguency with which Whistles and Keeaws Occurred Together and Separately With and Without a Known Disturbance as a Stimulus ........................ 56 Figure 16. Representative Sonogram of a Hiss .............. 58 Figure 17. Temporal Context With Which Two-act Sequences Occurred ......... ... . .............. . . . . . . . . . . . . . . . . i . . . 73 Figure 18. A Comparison Of Interaction Rates Between Colonies One and Two ................................... 75 Figure 19. Greeting and agonistic Interaction Rates per Dyad . ..... ...... ..... . .. . ..... ........ ................. 79 Figure 20. Variation i n Interaction Rates Among Months .... 80 Figure 21. Variation i n the Greeting Rate Throughout the Day . , 81 Figure 22. Home Ranges of the Two Adult Females (#12 and #15) in May 1974, and the Two Adult Males (#13 and #17) i n June 1974, on The Haley Lake Study Area ..... .................... .... ................ 83 Figure 23. Home Ranges of Four Female Marmots on the Haley Lake Study Area i n July 1974 84 Figure 24. Home Ranges of the Two Adult Males on the Haley Lake Study Area in August 1974 ......................... 85 Figure 25. Rates of Scent Marking by Adult Sales #4, #13, and #17 ........................................... 88 Figure 26. A Comparison of Greeting Rates Among Marmot Species ............ ... .....105 Figure 27. A Comparison of chasing Rates Among Marmot Species .................................... ............107 Figure 28. A Comparison of the Ratio of Greetings to Chases Among, Marmot Species ...................................108 LIST OF TABLES Table I. Habitat C h a r a c t e r i s t i c s of Vancouver Island Marmot Colonies ........................................ 13 Table I I . Climatic C h a r a c t e r i s t i c s of the Haley Lake Study Area and a Typical Parkland Subzone ( Location i n the Coast Mountains, afte r Brooke et a l • 1970 ........... 25 Table I I I . Age and sex Composition of Marmot Colonies One and Two ............................................35 Table IV. Causes of Whistling and Keeaw Sequences ........ 50 Table V. Dominance Matrices of Age and Sex Classes of Vancouver Island Marmots ...............................,64 Table VI. The Frequency Of Occurrence Of Each Social Behaviour Pattern Between Age-sex Classes 67 Table VII. Percentages of Social Behaviour Patterns per Age-sex Class 68 Table VIII. Comparisons of the Relative Frequency with which Social Behaviour Patterns Occurred Between Different Age-sex Classes ...................... 70 Table IX. Interaction Rate per Age-sex Class of Behavioural Acts per Thousand Hours .................... 76 Table X. Interaction Rate per Dyad per Behaviour per Thousand Hours ................ .................. ....... 77 Table XI. A Comparison of Vocalizations within the Marmota Caligata. Group ..................... .................... 92 Table XII. A L i s t Of Mammalian Species That Have Narrow Bandwidth Alarm C a l l s ..................................101 Table XIII. & Comparison of the Relative Growth Rates Of Y e a r l i n g Marmots ............................... ........110 X ACKNOWLEDGEMENTS I would l i k e to thank Dr. I. McT. Cowan for his supervision, encouragement, and patience throughout the course of t h i s project. I would also l i k e to acknowledge the help I received from: Ted Barsby, B r i t i s h Columbia Fish and W i l d l i f e Branch (Nanaimo), Dr. Fred Bunnell, Judith Donaldson, Jack Evans, Dr. N. R. L i l e y , MacMillan Bloedel (Chemainus D i v i s i o n ) , Dr. J . Mary Taylor, and Floy Z i t t e n . During t h i s study I was supported by a National Research Council of Canada Scholarship and Department of Zoology Teaching Assistantships. My research was financed by the World W i l d l i f e Fund (Canada). 1 INTRODUCTION SOCIOBIOLOGY AND BARASH'S HYPOTHESIS Recently, there has been a marked increase i n the number of investigators who are considering behaviour as a product of natural selection (Crook 1970, Barash 1974a S 1977, Alcock 1975, Brown 1975, Wilson 1975). Sociobiology (Wilson 1975) or socio-ecology (Crook 1970) i s the systematic study of the evolution of s o c i a l behaviour and s o c i a l organization i n r e l a t i o n to ecology, demography, and population genetics. This new approach to the study of animal behaviour has led to the formation of testable hypotheses concerning the adaptive si g n i f i c a n c e (contribution to f i t n e s s ) , of s o c i a l behaviour and s o c i a l organization. This study was an attempt to test one such hypothesis. Barash (1973b, 1974a) put forth a hypothesis to account for the evolution of marmot soc i e t i e s (Marjtota spp) . He showed that differences i n s o c i a l behaviour among marmot species were correlated with variations i n the environment, s p e c i f i c a l l y , the length of the vegetative growing season. Barash*s measure of growing season was the number of f r o s t - f r e e days in the absence of snow cover (Barash 1973b). Woodchucks (Warmota monax) inhabit environments of low elevation with long (150 day) growing seasons. They are s o l i t a r y and r e l a t i v e l y aggressive animals (Bronson 1964) and the young disperse from t h e i r natal burrows when weaned (Vos and G i l l e s p i e 1960). Olympic marmots (H. olyrnpus) inhabit high elevation alpine meadows that have a 2 very short (40 to 70 day) growing season. They l i v e in well integrated colonies and are s o c i a l l y tolerant, that i s , t h e i r s o c i a l behaviour i s characterized by a high freguency of greetings (Barash 1973b, 1971a). Olympic marmots do not mature u n t i l t h e i r fourth summer and disperse during t h e i r t h i r d . Barash's hypothesis i s that s o c i a l tolerance, as measured by the rate with which greetings are performed, i s inversely related to the length of the vegetative growing season. He reasoned that 1) shorter growing seasons r e s u l t i n lower growth rates in terms of the time reguired to reach adult size; 2) i t i s disadvantageous for subadult marmots to disperse when they are s t i l l "undersize", where s i z e i s measured as the weight of dispersing marmots r e l a t i v e to the weight of an adult of the same species; 3) in a l l marmot species, the minimum siz e for dispersers should be either the same proportion of the adult weight or dispersing marmots should be r e l a t i v e l y more mature i n species inhabiting increasingly severe environments; 4) aggression from adults causes the young to disperse; thus 5) the increase i n s o c i a l tolerance among marmots experiencing progressively shorter growing seasons may be due to the increasing necessity to i n h i b i t the dispersal of undersized animals. He also suggested that 1) i t i s important to have population size more cl o s e l y regulated as environmental severity increases because under severe conditions the habitat would be e a s i l y overgrazed, and there would be strong competition for food; and 2) i f marmot s o c i a l behaviour regulates population 3 size in a density dependent way; then 3) s o c i a l tolerance should increase i n severe environments because the operation of such a system of population regulation would require a closer physical proximity of colony members. Barash*s hypothesis i s testable in that i t predicts the degree of s o c i a l tolerance for any marmot species when the length of the vegetative growing season under which that species evolved i s known. Barash set out to test his own hypothesis by observing the s o c i a l behaviour of the yellow-bellied marmot (M. f/laviventris, Barash 1973a) , the hoary marmot (B. c a l i q a t a , Barash 1974b), and the European alpine marmot (M. marmota, Barash 1976b) . His observations of hoary and yellow-bellied marmots were consistent with his hypothesis, but alpine marmots engaged in s i g n i f i c a n t l y fewer greetings than were expected on the basis of the short growing season of i t s alpine habitat. Barash did not r e j e c t his hypothesis on the basis of this one inconsistent r e s u l t . He suggested that a blanket term such as s o c i a l i t y ( s o c i a l tolerance), may be inappropriate i n that i t obscures the differences between discrete parameters such as greeting and chasing, which may vary independently (Barash 1976b). Unfortunately, he does not go on to develop a revised hypothesis based on these discrete parameters. Armitage, Downhower, and Svendsen (1976) and Anderson, Armitage, and Hoffmann (1976) attempted to rej e c t Barash*s hypothesis because yellow-bellied marmots l i v i n g at high elevations grew fas t e r i n terms of grams per day than did yellow-bellied marmots l i v i n g at lower elevations. However, t h i s does not constitute a test of 4 Barash's hypothesis since his hypothesis i s based on the growth rate i n terms of the time taken to reach adult s i z e . The time taken to reach adult s i z e i s not related s o l e l y to the absolute weight gained per day, but i t i s also a function of the length of time each year during which marmots gain weight and the absolute weight of adult marmots of the species being considered. At the outset of this investigation there s t i l l remained two marmot species in North America which had never been studied, M. vancouverensis and M. broweri. The objective of t h i s study was to document the l i f e h i story and behaviour of Vancouver Island marmots and to provide another test of Barash's hypothesis of marmot s o c i a l i t y . VANCOUVER ISLAND MARMOTS The Vancouver Island marmot, Marmota yancouyerensis Swarth 1911, i s endemic to Vancouver Island, B r i t i s h Columbia. Li v i n g in small colonies on steeply sloping subalpine meadows, Vancouver Island marmots are active for only a few months each summer. To avoid the rigors of the mountain winters, marmots hibernate for about eight months of the year. Even then i t may be necessary for them to burrow out through many metres of snow in the spring. When I began t h i s study, t h i s was v i r t u a l l y a l l that was known about Vancouver Island marmots. This information was based on three very brief reports (Swarth 1912: 89-90, C a r l 1944, Hardy 1955: B61) which indicated that the natural history 5 of Vancouver Island marmots was s i m i l a r to other species of alpine marmots (e.g. Barash 1973b) and amenable to study using si m i l a r methods. Taxonomy and Evolutionary History Swarth described the Vancouver Island marmot as a new species i n 1911. The c r a n i a l and external c h a r a c t e r i s t i c s of M. -Vancouverensis are quite d i f f e r e n t from any other marmot species (Swarth 1911, Howell 1915). The most obvious c h a r a c t e r i s t i c s are: 1) the posterior border of the nasals which i s deeply V-shaped and, 2) the pelage which i s uniformly dark brown to black. However, the karyotype of M. vancouverensis i s very s i m i l a r to that of M. c a l i g a t a (Rausch and Rausch 1971). In 1915 Howell designated three groups of North American marmots based on morphological s i m i l a r i t i e s . He included M. vancouverensis i n the M. caligata group along with M« .caligata and M. olympus. In 1965, Rausch and Rausch considered PT. broweri to be a separate species within the i!« caligata group. This group appears to be a natural association for b i o l o g i c a l reasons as well as the purely morphological ones used by Howell. Species i n the M. c a l i g a t a group f i l l s i m i l a r e c o l o g i c a l niches and have si m i l a r behaviour (see discusion). Their ectoparasites also r e f l e c t the close evolutionary relationships within t h i s group. Fleas (Siphonaptera) collected from M. vancouverensis during this study were i d e n t i f i e d by Mr. G. P. Holland of Agriculture 6 Canada as Thrassis (Thrassis) spenceri spenceri Wagner. This subspecies of f l e a i s found only on marmots i n the M. c a l i j a t a group (Stark 1970). I c o l l e c t e d one t i c k (Acarina: Ixodidea) from M.vancouverensis. I am unaware of t i c k s having been collected from any other species i n the M. caligata group. Since t i c k s , l i k e f l e a s , tend to be host s p e c i f i c (Gregson 1956), i t was not surprising to discover that the specimen from M. vancouverensis may represent a new species i n the genus Ixodes (P. Zuk, Canadian Department of Agriculture, personal communication). Since only one subadult specimen i s available the species probably cannot be described. J» i§2co«verensis probably began to diverge from the ancestral M. ca l i g a t a stock af t e r crossing to Vancouver Island on temporary land connections and becoming isolated there. The fact that the following mammal species: musk ox Symbos cavifrons, mastodon Mammut americanum, mammoths Mammut imperator and Mammut cjolumbi, horse Eguus sp., and Bison sp. once l i v e d on Vancouver Island suggests that land connections with the mainland existed during the l a t e Pleistocene (Harington 1975). Harington (1975) believes that connections between Vancouver Island and the mainland existed on two occasions during the l a s t (Fraser/Wisconsin) g l a c i a t i o n and at least once during the penultimate (Illinoian) g l a c i a t i o n . The most recent connection with the mainland probably existed just p r i o r to the time of maximum development of continental ice during the Fraser g l a c i a t i o n about 20,000 years ago. At thi s time the sea l e v e l was depressed about 120 m below the present sea l e v e l and a 7 narrow corridor would have joined Port Angeles and V i c t o r i a (Fig 1). During the Peak of the I l l i n o i a n g l a c i a t i o n (roughly 100,000 years ago, Wright and Frey 196 5) sea l e v e l s were estimated to be 160 m below present l e v e l s and an even wider corridor would have joined Vancouver Island to the mainland. Marmots could also have crossed on the massive flo o d plain deposits that f i l l e d the whole S t r a i t of Georgia region (Fig 1) during the Olympia I n t e r g l a c i a t i o n about 30,000 years ago (date from F l i n t 1971) . Although l i t t l e i s known about the rate of animal speciation, Mayr (1963) has estimated that even a rapidly evolving island form would require a minimum of 100,000 years to achieve f u l l s p e c i f i c status. I f Mayr i s correct, i t i s most l i k e l y that marmots colonized Vancouver Island during or before the I l l i n o i a n g l a c i a t i o n rather than at either opportunity during the Fraser g l a c i a t i o n . The Vashon Stade of the Fraser g l a c i a t i o n covered most of Vancouver Island (Mathews, Fyles, and Nasmith 1970). Therefore, i f marmots crossed to Vancouver Island before t h i s period, there must have been either nunataks (high peaks and ridges emerging above the glaciers) or coastal refugia available on which the marmots could survive. Geological evidence indicates that both nunataks (Heusser 1960, Mathews et a l 1970, Muller unpublished) and coastal refugia (W. H. Mathews personal communication) existed. There are e x i s t i n g nunataks i n Alaska and the Yukon which are vegetated (Cooper 19 42, Heusser 1954) and support 8 a r c t i c ground s q u i r r e l (SoerraoDhilus undulatus) populations {Hurray and Hurray 1969) . Foster (1965) concluded that some species of mammals survived the l a s t g l a c i a t i o n on refugia on the Queen Charlotte Islands. Thus i t i s not d i f f i c u l t to imagine Vancouver Island marmots surviving the Vashon glaciacion on s i m i l a r refugia. Other zoological evidence also supports the refugia concept. No species of mammal other than vancouverensis presently e x i s t s exclusively i n the alpine-subalpine environment on Vancouver Island and no other mammal species on the i s l a n d has diverged to the point of being recognized as a d i s t i n c t species. In addition, white-tailed ptarmigan {Lagop_us leucurus) are also found i n alpine environments and a well d i f f e r e n t i a t e d race (L. leucurus sS^atalis) of t h i s r e l a t i v e l y implastic species i s confined to Vancouver Island (HcCabe and Cowan 1945). Other arguments for and against the nunatak survival hypothesis are discussed by Ives (1974). ..'..•!!•'-K-2Sil£2S2§ESS§i§- probably crossed to Vancouver Island via land connections that existed during the I l l i n o i a n g l a c i a l period and the species survived the g l a c i a l maxima of the Fraser g l a c i a t i o n on nunataks or coastal refugia or both. As the g l a c i e r s retreated free dispersal was probably made possible by the existence of alpine habitat at the edge of the retreating ice. As the forests closed i n below them Vancouver Island marmots would have gradually become i s o l a t e d on the mountain peaks that they now occupy. A s i m i l a r example of post g l a c i a l colonization of mountain tops has been postulated f o r the mountain hare (Leo us timid us) i n Europe (Hoffmann 1974) 10 METHODS STUDY AREAS I made most of my observations at two colonies situated about one kilometre apart on the southern end of Green Mountain, on Vancouver Island, B r i t i s h Columbia (numbers 1 and 2, Fig 1). The colony on the Haley Lake study area (colony number 1, Fig 1 and 2) has presumably been in continuous existence since i t s f i r s t discovery i n 1932 by K. Racey and I. McT. Cowan, since i t s c h a r a c t e r i s t i c s have not changed (I. McT. Cowan personal communication). I also made b r i e f observations on the other Green Mountain colonies (numbers 3, 4, and 5, Fig 1) and on Mt Washington, Mt Heather, and Buttler Peak (Fig 1, Table I ) . METHODS OF OBSERVATION I recorded observations on the behaviour of Vancouver Island marmots from 13 June u n t i l 16 September 1973 and from 30 A p r i l u n t i l 21 September 1974. The animals were observed from selected vantage points, without the use of a b l i n d , at distances of between 50 and 300 m. Observations were made through binoculars of 7x35 or 10x40 magnification, or spotting scopes of 15-60 or 20-45 power. Only observations occurring at least 15 minutes after my a r r i v a l were included i n my r e s u l t s — a time I judged to be s u f f i c i e n t for the marmots to habituate to my presence. Observations were recorded i n notebooks, on preconstructed t a l l y charts or with a tape recorder. 11 Figure 1. Locations of known Vancouver Island marmot colonies 1... Haley Lake s 2. Green Mt, co 3. Green Mt, co 4. Ski Club, co 5. Ski Club, co 6. Buttler Peak 7. Jordan Meado 8. Mt Hhymper ( 9. Heather Mt ( 10. Shaw Creek, 11. Mt McQuillam 12. Mt DeCosmos 13. Mt Moriarity tudy area (1976)* lony 2 (1974) lony 3 (1974) lony 4 (1974) lony 5 (1974) (197 4) ws (1930) 1971) 1974) headwaters (1944) (-Saunders; 1975) (?) (1971) 14. Golden Eagle Basin (1910) 15. King Solomon Basin (1910) 16. Mt Douqlas (1910) 17. Mt Arrowsmith (1938) 18. Cameron Lake (?) 19. Beaufort Range (1968) 20. Drink Mater Creek (1940) 21. Flower Ridge (?) 22. Golden Hinde (?) 23. Mt Albert-Edward (1970) 24. Mt Strata (1955) 25. Mt Washington (1974) 26. Coraox (1968; a dispersing marmot was collected near the c i t y ) * date pf the most recent confirmation of colony existence 11 a 32 km 12 Figure 2. Photographs of the Haley Lake study area ft. Photograph of the whole Haley Lake colony B. Close-up photograph of the part of the Haley Lake colony used most extensively by marmots {see also Fig 22, 23, and 24) 12a 13 Table I. Habitat c h a r a c t e r i s t i c s of Vancouver Island marmot colonies Colony Name North West Slope Aspect Elevation land Number Latitude Longitude (Degrees) (Meters) 1 Haley Lake 490 01* 1240 19* 33-63<»> SSE 1100- 1400 2 Green Mt 49° 0 1* 1240 19* 35< i> HSW 1 150- 1350 3 Green Mt 490 02* 1240 20* 33-45^2) SE 1500- 1600 4 Ski Club 49 0 03* 1240 20« 40<i> E 1550- 1600 5 Ski Club 490 03» 1240 20* 3 5-56<* > SSW 1550- 1600 6 Buttler Peak 490 00« 1240 20* 35-70<2> S 1500- 1550 7 Heather Mt 49 0 53' 1240 30* 33<2> S 1300- 1400 8 Mt Washington 490 40 • 1250 14* 2 0< 2> WSW 1500- 1600 degree of slope measured with a clinometer <f> degree of slope estimated 14 To obtain quantitative data on marmot behaviour I used a focal-animal sampling method (Altmann 1974). Focal-animal sampling consisted of selecting one individual and continuously recording i t s behaviour. Often more than one in d i v i d u a l could be observed at one time, but i t was never possible to observe a l l animals at a l l times (Ad libitum sampling, Altmann 1974). I was rarely faced with decisions on when to terminate observations on a f o c a l i n d i v i d u a l because f o c a l animals usually disappeared from view within a short time. Some observation periods were devoted to scan sampling; the recording of the behaviour and location of each animal every 10 minutes. ANALYSIS OF SOCIAL BEHAVIOUR DATA I considered a s o c i a l " i n t e r a c t i o n " to be an uninterrupted series of s o c i a l behaviour patterns or "acts" between two ind i v i d u a l s . Interactions were considered to be d i s t i n c t i f they were separated by an i n t e r v a l of more than one minute. I counted a s o c i a l behaviour pattern only once unless i t was separated by some other act other than t a i l r a i s i n g , since t a i l r a i s i n g always occurred at the same time as some other act. Interactions among three animals ( t r i a d i c interactions) were treated as a set of dyads (interactions between two animals). I a r b i t r a r i l y decided that a foc a l animal sampling period had to be greater than 15 minutes i n duration before I would include i t i n estimating the rates of interaction., I f e l t that 15 by only considering sampling periods that were longer than 15 minutes I would eliminate biasses inherent i n short samplinq periods. I calculated the rate, R, that any group of animals, i , (i . e . adult males), performed any s o c i a l behaviour, B, from the following formula: Si(B) Ri(B) = (1) SHi where Si(B) i s the number of a l l of the B acts involvinq or performed by group i during samplinq periods (S) , when individ u a l s in qroup i were the fo c a l i n d i v i d u a l s , and SHi i s the t o t a l time in hours (animal-hours) of sampling periods where indivi d u a l s i n group i were the f o c a l i n d i v i d u a l s . The rate, Ri(B), i s therefore an estimate of the number of B acts that animals in group i were involved in during each hour that they were active above ground and in plain view ( i . e . not per hour that I sat observing the colony). The units of Ri (B) are interactions per animal-hour. The rates of behaviour for individual marmots were obtained by considering the qroup as being composed of only one animal. The rate that group i performed act B with any s p e c i f i c group j , i s : 16 S i { B ) i : j + Sj{B)i:j IR(B)i: j = (2) (2) (SHi + SH j) where S i ( B ) i : j i s the sura of a l l B acts that occurred between i and j during sampling periods when the participant from group i was the f o c a l i n d i v i d u a l , . and there i s a 2 i n the denominator to correct for the fact that by watching each group independently I e f f e c t i v e l y counted each i n t e r a c t i o n twice. The units of IR(B)i:j are interactions per animal-hour, where animal-hours represents the number of hours that animals i n groups i and j were active above ground and in plain view. Some dyads never interacted during sampling periods, i n d i c a t i n g R i ( B ) i : j = 0, even though they may have interacted during observation periods which were of a shorter duration, thus i n d i c a t i n g that R i ( B ) i : j i s greater than zero. A non-zero estimate of R i ( B ) i : j can be calculated i f S i ( B ) i : j i s calculated rather than counted d i r e c t l y . The t o t a l number of B acts occurring between groups i and j over a l l observations regardless of t h e i r duration i s T { B ) i : j , and the t o t a l number of B acts occurring between group i and any other animal of known age and sex i s T(B) i . I f my observations of behavioural interactions were unbiassed then S i ( B ) i : j should be the same proportion of Si (B) as T ( B ) i : j i s of T(B)i. Thus, 17 S i ( B ) i : j T ( B ) i : j S i ( B ) T(B) i a n d S i (B) x T (B) i : j S i ( B ) i : j = {3) T{B) i S u b s t i t u t i n g e g u a t i o n (4) i n t o e q u a t i o n (3) y i e l d s : S i <B) x T (B) i : j S j ( B ) x T(B) i : j T (B) i T ( B ) j I R ( B ) i : j = (4a) (2) ( S H i + S H j ) E q u a t i o n 4a i s t h e f o r m u l a I u s e d t o c a l c u l a t e i n t e r a c t i o n r a t e s . I t was n e c e s s a r y t o be more e x p l i c i t i n t h e c a l c u l a t i o n o f i n t e r a c t i o n r a t e s f o r some s o c i a l b e h a v i o u r p a t t e r n s , s i n c e s o c i a l b e h a v i o u r p a t t e r n s c o u l d b e e i t h e r r e c i p r o c a l o r n o n -r e c i p r o c a l . I c o n s i d e r e d s o c i a l b e h a v i o u r p a t t e r n s t o be r e c i p r o c a l i f t h e a c t a p p e a r e d t o be a m u t u a l e x c h a n g e o f s i g n a l s b e t w e e n t h e i n t e r a c t a n t s ( e . g . g r e e t i n g , s e e a l s o s e c t i o n on S o c i a l B e h a v i o u r P a t t e r n s ) . N o n - r e c i p r o c a l b e h a v i o u r s were t h o s e i n w h i c h t h e i n t e r a c t a n t s a c t e d v e r y d i f f e r e n t l y f r o m e a c h o t h e r ( i . e . c h a s i n g ) . O n l y o n e i n t e r a c t i o n r a t e e s t i m a t e p e r d y a d was c a l c u l a t e d f o r e a c h r e c i p r o c a l a c t s i n c e t h e r a t e t h a t a n i m a l i g r e e t e d w i t h a n i m a l j was t h e same a s t h e r a t e t h a t j g r e e t e d w i t h i , i . e : H o w e v e r , i n t h e c a s e o f n o n - r e c i p r o c a l a c t s , t h e r a t e t h a t R i ( G ) i : j = R i ( G ) j : i a n d IB ( 6 ) 1 : j = IR (G) j : i . 18 a n i m a l i c h a s e d a n i m a l j i s not n e c e s s a r i l y e q u a l t o t h e r a t e a t w hich j c h a s e d i , i . e : R i ( C ) i : j * R i ( C ) j : i . Thus f o r e a c h n o n - r e c i p r o c a l b e h a v i o u r , two e s t i m a t e s o f i n t e r a c t i o n r a t e were c a l c u l a t e d f o r e a c h dyad, I R ( B ) i : j and I R ( B ) j : i . where S i { B ) x T ( B ) i : j S j ( B ) x T ( B ) i : j 4. T ( B ) i T ( B ) i I R ( B ) i : j = (2) ( S H i + SHj) and Si{B) x T(B) j : i S j ( B ) x T(B) j : i + T (B) i T ( B ) j IR(B) j : i = • (4b) (2) (SHi + SHj) The i n t e r a c t i o n r a t e s f o r a . s p e c i f i c d y a d c a n be added t o g e t t h e t o t a l i n t e r a c t i o n r a t e o r t o g e t t h e r a t e f o r any g r o u p o f b e h a v i o u r p a t t e r n s ( i . e . a l l a g o n i s t i c a c t s ) . T h e r e i s no s i m p l e r e l a t i o n s h i p between R i ( B ) and I R ( B ) i : j , when I R ( B ) i : j i s summed f o r a l l j ' s , b e c a u s e I R ( B ) i : j i s a f u n c t i o n o f SHj whereas R i ( B ) i s n o t . The f o r m u l a t h a t I have d e v e l o p e d f o r c a l c u l a t i n g i n t e r a c t i o n r a t e s y i e l d t h e same r e s u l t s as t h o s e e q u a t i o n s used by A r m i t a g e (1976a) f o r some d y a d s . However, A r m i t a g e c a l c u l a t e s mean r a t e s i n c o r r e c t l y by a d d i n g r a t e s i n "any c o m b i n a t i o n d e s i r e d " ( A r m i t a g e 1976a). 19 Unless otherwise indicated, a significance level of 0.05 was used for a l l s t a t i s t i c a l t e s t s . TRAPPING AND MARKING In 1974 marmots were captured in 2 5x30x80 cm or 22x22x63 cm l i v e traps manufactured by Tomahawk Live Traps Co., Tomahawk, Wisconsin, 0. S, A. Baits used were peanut butter, peanuts and the leaves and flowers of preferred species of food plants when these were available (Appendix I ) . Marmots were transferred from the traps to a handling cone s i m i l a r to the one i l l u s t r a t e d by Taber and Cowan (1971). No t r a n g u i l i z a t i o n was necessary. Animals were measured using a f l e x i b l e s t e e l millimetre tape while being held as nearly as possible i n an extended position. I recorded the sex, and weighed the marmot with a 12 Kg spring balance (Pesola Scales, Basle, Switzerland) that could be read to the nearest 100 g. The occurrence of ectoparasites and a description of the molt was also recorded. Each animal was marked by attaching a single numbered rabbit ear tag (style #4-1538) supplied by the National Band and Tag Co., Newport, Kentucky, U. S. A. With each tag one or two coloured p l a s t i c markers of Dymo embossing tape (Dymo of Canada Ltd., Missisauga, Ontario), approximately 1x2 cm, were applied. Ear tags were placed as far from the margin of the ear as possible before the thickness of the ear became l i m i t i n g . Tags were occasionally l o s t by other animals bi t i n g the coloured markers and ripping the whole tag from the ear. Tags were most 20 frequently l o s t i n the traps or during handling. Most animals were tagged at least twice during t h i s study. In an attempt to permanently mark animals I t r i e d the freeze branding techniques described by F a r r e l l , Roger, and Winward (1966), Hadow (1972) and Ch u r c h i l l and Coburn (unpublished). The desired r e s u l t of freeze branding i s a regrowth of white hair i n the shape of the brand which can be recognized at a distance. , This result requires that the melanocytes be destroyed but not the hair f o l l i c l e . In previous laboratory studies, the regrowth of white hair occurred in 3 to 6 weeks and was retained through subseguent molts ( F a r r e l l et a l 1966, Taylor 1969, Hadow 1972, C h u r c h i l l and Coburn unpublished, Lazarus and Rowe 1975). I used two brass branding "irons:" one a 1x5 cm rectangle, and the other a c i r c l e , 3 cm i n outside diameter and 1.6 cm i n inside diameter. Each brand therefore had a surface area of 5 cm2. Each animal was marked with a unique combination of two brands by varying the brand orrientation and position on the body. Brands were applied to a shaved area on the body for exactly 30 seconds. 21 MEASUREMENT OF MICROCLIMATE In 1973 daily maximum and minimum temperatures sere recorded. In 1974 a l l of the following measurements were obtained. Daily r a i n f a l l was recorded with a Tru-Check Rain Gauge (Tru-Check Inc., Albert Lee Minnesota). A continuous record of temperature and humidity was obtained from 12 June u n t i l 3 November using a 31 day Casella thermohygrograph. The thermohygrograph was enclosed i n a Stevenson Screen which was situated on a small eminence in the centre of colony number one. Additional weather information was obtained from instruments operated by the Secretariat for the Environment Land Use Committee of the Province of B r i t i s h Columbia. These instruments were located approximately 300 m below colonies 5 and 6. VOCALIZATIONS I recorded vocalizations of both trapped and free ranging marmots with a Uher 4000IC Report tape recorder. The microphone used was either an ElectroVoice 64 4 Sound Spot d i r e c t i o n a l microphone or a Sony Dynamic M136 microphone. The recording speed was always 19.05 cm per s. Sonograms were made from these recordings on a Kay Co. Missilyzer model 675. Recordings of both trapped and free ranging marmots were used i n the cal c u l a t i o n s of whistle length and freguency. A l l other data were from free ranging marmots only. 22 RESULTS HABITAT CHARACTERISTICS Vancouver Island aariots l i v e in subalpine habitats that are characterized by steep c l i f f s , talus debris, and open meadows that are usually oriented south of the east-west l i n e (Table I, Fig 2 and 3). Below the talus, the slope becomes less steep, the substrate becomes more stable, and herbaceous plant communities develop. Where the slope i s steep enough, avalanches and snow creep i n h i b i t the establishment of trees. Evidence of these forces can be seen in the form of uprooted saplings and the d i s t i n c t basal crook in a l l established trees (Fig 3A) . , A l l of the marmot colonies that I v i s i t e d had plant communites that were c h a r a c t e r i s t i c of the Parkland Subzone of the Subalpine Mountain Hemlock Zone of B r i t i s h Columbia (Brooke, Peterson, and Krajina 1970). Within the Parkland Subzone Brooke et a l (1970) describe eight plant associations. Plants found on the Haley Lake study area such as mountain hemlock Tsuga fflgrtengiaS5' yellow cedar Chamaecyparis nootkatensis. blue-leaf huckleberry Vaccinium delicjosum, partridgefoot Leutkea Bgctinata, and mountain daisy Eriaeron peregrinus (see also Appendix I) were c h a r a c t e r i s t i c of the Nano-tsugetum mertensianae association, subassociation nano-tsugetum mertensianae. Other plant associations were also present on the Haley Lake study area and on other colonies. Three colonies, Mt 23 Figure 3. Photographs of Vancouver Island marmot habitat A. The Haley Lake colony i l l u s t r a t i n g a t y p i c a l open meadow habitat B. The Mt Washington colony i l l u s t r a t i n g the dense cover of white rhododendron and alpine f i r 23a B 24 Washington, Heather Mt> and Central Green Mt (colony 3) d i f f e r e d from the others i n that they were not below c l i f f s . Open meadows were c h a r a c t e r i s t i c of a l l marmot colonies except Mt Washington. Marmot burrows on Mt Washington were located i n an area with numerous small, 1-5 m, trees, mainly alpine f i r Abies lasiocarpa. The area was also covered by a dense growth of white rhododendron Rhododendron albiflppum. and Vaccinium sp. (Fig 3B) . The: climate of the Subalpine Mountain Hemlock Zone i s characterized by cool short summers and wet winters with considerable p r e c i p i t a t i o n f a l l i n g as snow (Table I I ) . The length of the growing season i s d i f f i c u l t to determine i n subalpine areas since minimum a i r temperatures are often well above freezing while snow s t i l l persists on the s i t e due to the great accumulations during the winter (Brooke et a l 1970 and Table I I ) . On the Haley Lake study area i n 1974, the number of f r o s t - f r e e days was above average at 135 days (Table I I ) , but the e f f e c t i v e growing season was about two weeks less on much of the colony (Table II) as a re s u l t of a persistent snowpack. However, the snow pack was not evenly di s t r i b u t e d . The c l i f f s above the colony were too steep to accumulate much snow and therefore became snow free much e a l i e r than the rest of the colony. The c l i f f s on the other colonies were also the f i r s t areas to become snow free. A small part of colony 2 was free of snow very early i n 1974, apparently having been kept r e l a t i v e l y snow free by the prevailing wind. Marmots foraged on these early snow free areas u n t i l the snow melted from the main part 25 Table I I . Climatic c h a r a c t e r i s t i c s of the Haley Lake study area and a t y p i c a l Parkland Subzone location i n the Coast Mountains, after Brooke et al 1970 , Climatic Baley Lake Year Brooke et a l Chara c t e r i s t i c s Study Area 1970 annual p r e c i p i t a t i o n i n cm and percent snow r a i n f a l l from June through September (cm) mean temperature July through September (°C) date of the l a s t f r o s t i n the spring number of f r o s t - f r e e days maximum accumulation of snow (cm) l a s t accumulation of show i n the spring 25 1974 13.0 1973 15.5 1974 13-20 June 1973 3 June 1974 115 1973 135 1974 approximately 300 1974 early June 1973 la t e June 1974 285 35% 50 12.0 end of May 114 370 l a t e June 26 of the colony. On the Haley Lake study area i n 1974 marmots had to forage on the c l i f f s for 6 weeks afte r emergence i n the spring. Steep slopes are c h a r a c t e r i s t i c of Vancouver Island marmot habitat because they are susceptible to avalanches. Avalanches provide suitable habitat 1) by maintaining herbaceous communities through the i n h i b i t i o n of tree growth, and 2) by reducing the accumulation of snow, r e l a t i v e to f l a t t e r areas, which i n turn r e s u l t s i n the meadow becoming snow free e a r l i e r in the spring, thus increasing the e f f e c t i v e growing season. The average area of 8 marmot colonies was about two hectares (range 0.5 to 4.0). PHYSICAL CHARACTERISTICS I recognized the following age classes of Vancouver Island marmots: infants, yearlings, two-year-olds, and adults. I d e n t i f i c a t i o n of age classes was based on weights (Fig 4). I did not trap any infants during t h i s study, but infants were e a s i l y distinguished because of t h e i r small size (Fig 4). My sample s i z e was too small to compare the differences between the weights of male and female yearlings, but yearlings as a group weighed less than two-year-olds (F=123.13; d.f. = 1,16; p<0.001). Some females had weights that were s i g n i f i c a n t l y greater than female yearlings (F=65.6; d.f.-1,13; p<0.001) but 27 Figure 4. Seasonal changes in the weights of Vancouver Island marmots The growth rate equations are as follows: Adults, sexes combined y = 0.0245x • 2.466 Two-year-old females y = 0.0259x + 1.516 Yearlings, sexes combined y = 0.0230x + 0.584 where y i s the weight i n kilograms and x i s the number of days since spring emergence (30 April) * known aged two-year-old 2 infant female; data from a specimen collected from Mt Washington i n 1965, University of Alaska specimen #28754 2?a 28 s i g n i f i c a n t l y less than the largest females (F=18.7; d.f.=1,22; P<0.001). I assumed that these animals were two-year-olds. In 1975, I captured one female known to be two years old. She weighed s l i g h t l y less than the average two-year-weight predicted from the'growth rate equation (Fig 4). This observation i s consistent with the assumption that two-year-olds weigh less than adults. Two-year-olds can be recognized by weight i n M. c a l i g a t a (Barash 1974b) and M. olympus (Barash 1973b). A l l males that were non-yearlings had similar weights. The absence of any males recognizable as two-year-olds could be the resu l t of the absence of any two-year-old males on my study areas or the growth of males being such that they reach their adult (heaviest) weight as two-year-olds. In M. olympus* two-year-old males are s t i l l distinguishable from adult males on the basis of weight (Barash 1973b). I assumed that the same would be true for Vancouver Island marmots and that I had no two-year-old males on my study areas. The term adult, then, r e f e r s to animals that are considered to be at least three years old. Adult males weighed s i g n i f i c a n t l y more than adult females (Fig 5; F=6.87 d.f. = 1,28 p=0.014). Growth rate appears to be l i n e a r for a l l sex and age classes throughout the summer (Fig 4 and 5). Analysis of covariance indicated that the rate of weight gain did not d i f f e r s i g n i f i c a n t l y among sex and age classes. Barash (1973b) used the change i n M. olympus- tooth colour 29 Figure 5. Seasonal changes i n the weights of adult marmots The growth rate equations are as follows: adult males y = 0. 0210x «• 2. 56 1 adult females y = 0.0263x • 2.489 where y i s the weight in kilograms and x i s the number of days since spring emergence {30 April) . 29a 3 V B-1 7-1 • A Y S AFTER SPRING EKERGEISCE C30 A P R I L ) 30 (from d u l l w h i t e t o d a r k o r a n g e ) as an a i d i n s e p a r a t i n g age c l a s s e s . T h i s was n o t p o s s i b l e i n V a n c o u v e r I s l a n d marmots s i n c e t h e t o o t h c o l o u r o f a l l V a n c o u v e r I s l a n d marmots was d u l l w h i t e . The f r e s h p e l a g e o f M. v a n c o u v e r e n s i s i s b l a c k o r v e r y d a r k brown. T h i s c o l o u r i s u n i f o r m o v e r most o f t h e body e x c e p t f o r a c o n s p i c u o u s p a t c h o f w h i t e f u r a r o u n d t h e n o s e and mouth, a s m a l l w h i t e mark on t h e f o r e h e a d , and some w h i t e s t r e a k i n g on t h e b r e a s t and abdomen. The c o l o u r o f t h e f r e s h p e l a g e g r a d u a l l y f a d e s o v e r t h e summer and f o l l o w i n g s p r i n g t o a l i g h t c i n n a m o n brown. V a n c o u v e r I s l a n d marmots molt o n l y once p e r y e a r . The p r o g r e s s i o n o f t h e m o l t was e a s i l y o b s e r v e d i n t h e f i e l d b e c a u s e t h e f r e s h p e l a g e c o n t r a s t e d s o s h a r p l y w i t h t h e o l d f a d e d f u r . . A n i m a l s f i r s t showed s i g n s o f m o l t i n g i n mid-J u l y . The new f u r emerged f i r s t on t h e rump, o r on t h e f o r e l e g s and s h o u l d e r s . The emergence o f f r e s h p e l a g e o v e r t h e r e s t o f t h e body was e x t r e m e l y v a r i a b l e . T h e l a s t a r e a s t o molt were u s u a l l y t h e back o f t h e head, t h e t a i l , and t h e rump. Some a n i m a l s d i d n o t a p p e a r t o c o m p l e t e t h e i r molt by t h e t i m e t h e y h i b e r n a t e d i n l a t e September; n o n e t h e l e s s , m o l t i n g f i n i s h e d a t t h i s t i m e . I n c o m p l e t e l y m o l t e d a n i m a l s emerged f r o m h i b e r n a t i o n ! t h e n e x t s p r i n g a t t h e same s t a g e o f m o l t as t h e y were a t t h e p r e v i o u s f a l l , and no s u b s e q u e n t change o c c u r r e d u n t i l t h e n e x t J u l y . I f an i n d i v i d u a l had n o t c o m p l e t e d i t s m o l t f r o m t h e p r e v i o u s y e a r , t h e molt began f i r s t i n t h e a r e a s h a v i n g t h e 31 oldest pelage. Young animals emerged from the burrows with black fur. They did not appear to molt during t h e i r f i r s t summer but molting would be d i f f i c u l t to detect because of the absence of any colour change. I could not detect any other differences i n molting c h a r a c t e r i s t i c s among di f f e r e n t age and sex classes. Davis (1966) noted that M. mpnax also had a v a r i a b i l i t y in the progression of the molt, although i t always began on the rump. He also observed that some animals did not complete th e i r molt each year. One molt per year appears to be the rule among marmots, W. raonax (Hamilton 1934, Davis 1966), vancouverensis (this study), M. f l a v i v e n t r i s (Armitage 1974), and M. olympus (Walker 1964) , although t h i s was disputed by Barash (1973b) for A l l freeze brands were applied on either the 23, 24, or 25 July 1974. This was the e a r l i e s t date possible because a labour s t r i k e prevented the production of dry ice u n t i l this time. At the time of appl i c a t i o n , fresh pelage had already begun to grow over much of the body. Four weeks after branding no regrowth of fur had occurred. The branded areas were bare, the club hairs and a layer of skin having been sloughed off. Seven weeks after branding, the brands had s t i l l not grown in completely. The fur which was i n the process of growing in was either black, the normal colour of fresh pelage, white, as expected, or rar e l y , black with white t i p s . No brands were v i s i b l e at a distance as white fur in 1974. In June 1975 the brands had s t i l l not 32 regrown on the one i n d i v i d u a l for which detailed observations were possible. No animals displayed white brand markings which were v i s i b l e from a distance at th i s time. In August 1975, the brands on one i n d i v i d u a l had regrown with normal coloured pigmentation. However, three marmots had brands that were i d e n t i f i a b l e at a distance of about 100 m when viewed through a 15-60 power spotting scope. On close observation I found that these brands were only sparsely covered with white fur. Since on Vancouver Island marmots the white fur did not become v i s i b l e u n t i l the next molt a f t e r branding, over one year l a t e r , freeze branding was useless as a marking technigue f o r t h i s study. However, i n the future the e f f i c a c y of freeze branding might be increased i n two ways. One, branding could be done e a r l i e r i n the summer to minimize the time between branding and the beginning of the molt. This could re s u l t in brands becoming v i s i b l e in three to six weeks as in the laboratory studies (Hadow 1972, C h u r c h i l l and Coburn unpublished, Lazarus and Rowe 1975). A second improvement that could be made i s the length of time during which the brand was applied. The application time i s very important to achieve good r e s u l t s ( F a r r e l l et a l 1966, Hadow 1972, C h u r c h i l l and Coburn unpublished). Free ranging marmots may reguire a d i f f e r e n t branding time than was predicted from experiments with Abort*s s g u i r r e l s (Sciurus aberti) and fox s q u i r r e l s (Sciurus niger) (Hadow 1973) and Columbian ground s g u i r r e l s {Spermophilus £2l]iSfei§llSS; C h u r c h i l l and Coburn unpublished) . It i s not clear from my re s u l t s whether the branding time that I used was too 33 short or too long ( i f e i t h e r ) . Since the brands took a long time to regrow and normal pigmentation sometimes developed, the application time appeared to be too long (Hadow 1972). However, the r e s u l t i n g intermediate pigmentation i n some animals indicated that the application time was too short (Churchill and Coburn unpublished). Lazarus and Bowe (1975) used a commercially available pressurized refrigerant as a freeze branding agent. Their re s u l t s are extremely encouraging and t h e i r technigue appears to be much more e f f i c i e n t than using dry i c e , es p e c i a l l y under f i e l d conditions. A pressurized refrigerant available i n Canada s i m i l a r to the one used by Lazarus and Bowe (197 5) i s Can.O.Gas Refrigerant 12, V i r g i n i a Chemicals Inc., Portsmouth, Va. U.S.A. COLONY COMPOSITION I considered a colony to be a group of animals that was is o l a t e d both geographically and s o c i a l l y from other such l o c a l i t i e s . I knew the exact population size for two colonies, colonies one and two on Green Mountain (Fig 1). These r e s u l t s are presented in Table I I I . Since I did not trap animals i n 1973, I was only able to recognize infants and older animals and was unable to determine t h e i r sex. However, I knew the status of most animals i n 1974. The average colony size in June, before the infants emerged above ground, was 8.3 animals (n=4. Table I I I ) . A l l the other colonies that I observed appeared to be of a si m i l a r s i z e . The average s i z e of f i v e l i t t e r s was 3.0 34 ±.32 (SE) infants (n=5) . Movement of marmots between colonies was very rare. , There were only four occasions, that I knew of, when marmots moved between colonies on Green Mountain., I saw tracks of marmots in the snow between colonies one and two on 20 May 1974 and most of the way from colony one to colony three on 11 June 1974. One adult male was observed on colony one u n t i l 2 June 1974. On 12 June he was seen on colony two where he remained f o r the duration of the summer., An adult male immigrated to colony one from an unknown location on, or shortly before, 25 June 1974. He remained there for the duration of the summer. Colony one therefore was occupied by three d i f f e r e n t adult males i n June 197,4; however, there were only two present at any one time (Table I I I ) . I was unable to determine the sex of three marmots when I f i r s t captured them., They were rarely seen subsequently and were never recaptured; therefore, t h e i r sex was never determined. , ACTIVITY PATTERNS The general pattern of Vancouver Island marmot a c t i v i t y i s guite s i m i l a r to that reported for other marmot species (Armitage 1962, Gray 1967, Barash 1973b, Hayes 1976). Vancouver Island marmots have an annual schedule made up of a summer a c t i v i t y period of 4 to 5 months and a winter hibernation period of 7 "to 8 months. In 1974 marmots were active on 30 A p r i l , the f i r s t day that I v i s i t e d the Haley Lake study area. Tracks i n 35 T a b l e I I I . A q e a n d s e x c o m p o s i t i o n o f m a r m o t c o l o n i e s o n e a r d t w o N u m b e r o f M a r m o t s c f E a c h A q e a n d S e x C c l c n y Y e a r M o n t h AM AP A? 2F 2? YH Y F Y ? I ? TOTAL 1973 J u n e 11 11 J u l y 11 6 17 A u q 8 4 12 S e p t . 8 4 12 1974 H a y 2 2 1 1 1 2 1 11 J u n e 2 2 1 1 1 1 2 1 11 J u l y 2 2 1 1 1 2 1 10 A u g 2 2 1 1 2 8 S e p t 2 2 1 1 6 1973 J u n e 6 6 J u l y 4 2 6 Aug 4 2 6 197 4 May 1 1 1 1 4 J u n e 1 1 1 1 1 5 J u l y 1 1 1 1 4 A u g 1 1 1 1 4 A = A d u l t ? = S e x u n k n o w n I = I n f a n t • 1 - F e m a l e M = M a l e 2 = T w o - y e a r - o l d Y = Y e a r l i n g 36 the snow indicated that marmots had probably not been active for very long and a l l in d i v i d u a l s had not necessarily ended their hibernation at t h i s time. There was no sign that any animals had yet emerged when I v i s i t e d the Haley Lake study area on 17 A p r i l 1975. I l a s t saw marmots on 16 September 1973 and 21 September 1974. I observed various colonies for many hours, on several ;days after these dates. Naturalists recorded seeing marmots on the Haley Lake study area on 30 September and 1 October 1972 (Bob Morris and Ted Barsby personal communication). Thus I considered early May to be a reasonable estimate of spring emergence with most ind i v i d u a l s in hibernation again by mid-September. I did not notice differences among age-sex classes with respect to the time of spring emergence or f a l l hibernation but I have few observations from these periods. It would be guite unusual i f Vancouver Island marmots did not have age s p e c i f i c emergence and hibernation times since t h i s i s c h a r a c t e r i s t i c of a l l other marmot and ground s g u i r r e l species that I know of (e.g. Vos and G i l l e s p i e 1960, Armitage 1962, Iverson and Turner 1972, Yeaton 1972, Barash 1973b). Both l i t t e r s of infants that were born on the Haley Lake study area i n 1973 emerged from the i r burrows f o r the f i r s t time on 11 July. Swarth (1912) noted that i n the Mt Douglas area no l i t t e r s had emerged by the t h i r d week in July 1911. I compiled a c t i v i t y budgets for each i n d i v i d u a l i n terms of the percent of the t o t a l observation time that a marmot spent 37 resting (lying or s i t t i n g outside of the burrow), feeding, i n the burrow, moving (without feeding), engaging i n s o c i a l behaviour, and grass c o l l e c t i n g . A c t i v i t y budgets f o r the months of May through September are presented i n Figures 6 to 10. Only the data for resting, feeding, and i n burrow time are presented since these three behaviours accounted for more than 93% of the time budgets of a l l animals i n each month.. The data were combined f o r a l l individuals since there were no s i g n i f i c a n t differences among age-sex classes with respect to these three a c t i v i t y patterns. The daily pattern of a c t i v i t y varied with the time of year. A midday a c t i v i t y l u l l was not apparent i n May and September. In June, July, and August marmots were much more active i n the mornings and evenings than at midday. The obvious decline in midday a c t i v i t y i n July and August seemed to be the res u l t of three f a c t o r s ; temperature, an inherent circadian rhythm, and a general decline in the amount of time spent feeding as the summer progressed. ....... I divided the day into three periods, morning, midday, and evening, based on the duration of a marmot day. These periods varied s l i g h t l y between months but the midday period was usually between 1100 and 1600 hours. The maximum daily temperature occurred during the midday period except on a few unusually cool days. The percent of time spent feeding at midday dropped o f f dramatically when the maximum daily temperature exceeded 20°C. 38 Maximum Daily Temperature (°C) Percent During of Time Spent Feeding the Midday Period 0 - 14.9 23 15 - 19.9 22 20 - 24.9 6 25 - 29.9 0 also, the time spent i n the burrow at midday was d i r e c t l y correlated with the maximum da i l y temperature {r=0.69, p<0.01). Temperatures over 20°C were much more common in July and august than i n the other months. When the maximum temperature was less than 20°C marmots s t i l l fed sigi.fica.ntly l e s s during midday than they did during the morning and evening periods {data from morning and evening periods were lumped and tested against the midday period; t=2.90, df=63, p<0.01). Thus there was a bimodal pattern of d a i l y feeding a c t i v i t y which was accentuated by high temperatures. The t o t a l time spent feeding per day decreased throughout the summer. This trend was probably the r e s u l t of an increase in both food guality and guantity over the summer. As the summer progressed vegetation guantity increased as the snow melted and vegetation guality increased as more species came into flower. Flowers are more nutritious than vegetative parts (Svoboda 1972) and are selected by marmots when available {Appendix I ) . as the time required to obtain s u f f i c i e n t food decreased, there would be l e s s pressure to feed at midday, further accentuatinq the a c t i v i t y l u l l s i n July and august. The 39 Figure 6. A c t i v i t y budgets for May, a l l animals combined 65 animal-hours of observation MAY ACTIVITY CALL ANIMALS) 39a 100. 90-. . 80. . . 70. . . 60. . . 50-. -40.. . 30- . . ao... 10.. . 0. 10- 11 > 1H> 13- 14- IS- !£• 17- 18- 19. BO-HOUR DF THE DAY 10O-90- . . 80. . . 70- . . GO... 50v. 40-.. 30. . . 30' 10 0 10. i l . 12- 13. 14. 15. IB- 17- IB- IS- BO HOUR OF THE DAY 100. 90. 80-70-GO-50- , 40- . 30-. 50.. 10-. 0.. 10- 11- 12- 13- 14. IS. 17. IS. 19- 50. H X R OF THE DAY 40 F i g u r e 7. A c t i v i t y budgets f o r June, a l l animals combined 175 animal-hours of o b s e r v a t i o n 40a JLNE ACTIVITY CALL ANIMALS) 100. 90... BO... 70... 60... 50... 40... 30-.. 30... 10... 0 - . _ 4- 10- 11. IS- 13. 14- 15- 16- 17. IB' 19- SO- HI. 33. SS-I-CUR CF THE DAY 100-90. 80-70-eo-50-40-30. so. 10. o. 10. 11. 15 < 13- 14. 15. 16> 17. 18. 19- 50. 51- HE- S3-HOUR OF THE OAY 100-. 90-. 80-. 70-. 60-. 50-. 40". SO-SO-. 10-0-9. 10- 11- IS" 13- 14- 15> 1G- IB- 19. SO- E l . E2« S3> HOUR DF THE DAY 41 Figure 8. A c t i v i t y budgets for July, a l l animals combined 202 animal-hours of observation 4la JULY ACTIVITY CALL ANIMALS) 100-90. . . BO... 70:. GO... 50. . . AO:. 30. . . 50. . . 1X3... 0 . ._ 9- 10. ! ! • 15- 13- 14- 15- 16- 17- IS' 19- SO- 51« SH- B3> HOUR OF THE DAY 100-90- . . 90- . . 70- . . 60 . . . 50. . . 40« . . 30- . . 50- . . 10- . . 0 . . . 1 1 1 6* 7. 6> 10. i l ' ' lS . 13 • 14. 15- IB- 17. IB' 19- SO- 51 • 55' S3-HOUR OF THE DAY 100-90 ' . . BO-.. 70. . . GO 50. . . 40. . . 30. . . SO-.. 10-0 4. 5- 10. 11. 12- 13- 14- 15- IB- 17- IB' 19- 50- 51- 25- S3-HOUR CF THE DAY 42 Figure 9. A c t i v i t y budgets for August, a l l animals combined 114 animal-hours of observation 42a AIGLET ACTIVITY CALL ANIMALS) 100. 30-.. 80-.. 70... BO. . 50... 40. . 30-.. SO... IX)... 0-._ 10. 11. IS- 13- 14- 15- IB- IB- 13- SO- 51- SB- 53-HOJR OF THE OAY 100-90... 90-.. 70- . SO... 50-.. 40- . 30-.. SO-.. 10-.. 10- 11. IS- 13- 14. 15- 15- 17- IB- 19- SO. HI. S3-HOUR DF THE DAY 100-90-SO-70-60-. 50-40-30-HO-10-0-. 10. 11. IH. 13- 14- 15> IE- 17- 18- 19- HO- HI. EE- S3-HXR CF THE DAY 43 Fi g u r e 10. A c t i v i t y budgets f o r September, a l l animals combined 70 animal-hours of o b s e r v a t i o n 43a SEPTEMBER ACTIVITY (ALL ANIMALS) 100. 90... BO... 70-.. 60... 50... 40-.. 30-.. eo... 10... 0. 9- 10- 11. 12. 13- 14. 15- 16- 17- IB' 19. 20. 21- 22 HOUR OF THE DAY 100 90-.. SO... 70-.. BO-.. 50 40-.. 30... 20... 10... 0 10. 11. 12- 13- 14. IS. 16- 17. IB- 19. 20. 2 1 . E 2 HOUR OF THE DAY 100-90-.. BO... 70... BO. 50... 40... 30... 20... 10... 0. R- 10- 11- 12- 13- 14. 15- I B - 17. IB- 13- BO. 21. 22. HOUR OF THE DAY 44 absence of a midday a c t i v i t y l u l l i n September may be an e f f e c t of short day length or metabolic changes of upcoming hibernation on the basic circadian rhythm. I saw marmots c o l l e c t i n g the brown dead stems of grasses and sedges and taking them i n t o t h e i r burrows on 152 occasions. I assumed that t h i s material was used for bedding since I never saw marmots eating i t above ground. The frequency of grass c o l l e c t i n g decreased through the summer but there was a s l i g h t increase i n September just prior to hibernation. Adult females collected more often than the other age-sex classes. I observed marmots c o l l e c t i n g grass at a l l times of day, but i t occurred most often at the very end of the day, just before they entered the burrow for the night. VOCALIZATIONS Whistles The sound most frequently produced by Vancouver Island marmots i s a loud piercing "whistle" which originates i n the vocal chords. The dominant frequency of a whistle i s the f i r s t harmonic or fundamental frequency. The f i r s t harmonic occurs at 2910±33 Hz (n=36). Whistles also possess a second harmonic at about 5700 Hz and a t h i r d harmonic at about 8400 Hz. Both of these harmonics contain much less energy than does the fundamental frequency (Fig 11 and 12). The i n t e n s i t y of the sound did not vary appreciably throughout the duration of the 45 Figure 11. Representative sonogram of a short whistle Recording speed: 19.05 cm per s Playback speed: 4.76 cm per s Narrow bandwidth f i l t e r eoooo. 1BOOO-. 1GOOO-. 14O0O-. iaooo._ 10000.. TIME ( S E T O N D S ) 46 F i g u r e 12. R e p r e s e n t a t i v e sonogram of a l o n g w h i s t l e Recording speed: 19 . 05 cm per s Playback speed: 9 . 3 5 cm per s Wide bandwidth f i l t e r 46a 47 whistle. I subjectively c l a s s i f i e d most marmot whistles that I heard i n the f i e l d as being either "long", "medium", or "short" i n duration. I then used sonograms to determine the duration of each of these whistle types. The results were: whistle mean c l a s s i f i c a t i o n duration (s) range (s) SE n long 0.57 0.27 - 0.84 0.052 14 medium 0.26 0.23 - 0.29 0.009 7 short 0.20 0.12 - 0.26 0.009 17 The i n a b i l i t y to d istinguish medium whistles was not serious since most whistles that I heard i n the f i e l d were c l e a r l y either long or short. Medium whistles were therefore omitted from subsequent analysis except where noted. There was no s i g n i f i c a n t difference between the whistles of trapped and free ranging marmots with respect to duration (t= 1. 50) or freguency (t=l.05). ft whistling sequence was considered to be any whistle or group of whistles that were les s than one minute apart. Most whistling sequences consisted of a single whistle (Fig 13). The longest sequence of long whistles was 50 whistles i n 9 minutes. One sequence of short whistles lasted 28 minutes and contained about 400 whistles. The i n t e r v a l between whistles varied with the length of the whistle. Long whistles had a mean i n t e r -whistle i n t e r v a l of 17.9 ±1.59 s and short whistles averaged 2.9 48 F i g u r e 13. ft comparison of the l e n g t h of long and shor t w h i s t l i n g sequences BO. 1 2 3 4 5 >5 NUMEER CF WHISTLES IN A WHISTLING SEQLENCE 4.9 ±0.08 s between whistles. Marmot whistles serve primarily a warning function (Armitage 1962, Waring 1966, Barash 1973b, 1975).. Vancouver Island marmots always whistled when cougars (Felis concolor), black bears (irsus americanus) golden eagles (Aquila chrysaetos), bald eagles (Haliaetus leucocephalus), or red-t a i l e d hawks (Buteo jamaicensis) were detected. These species are probably the major predators of Vancouver Island marmots. On one occasion I saw a golden eagle make an unsuccessful attack on a marmot. On another, I am guite sure that two cougars caught a marmot at a spot where trees p a r t i a l l y obscured my view. A l l of these predators have been reported to prey on other marmot species (cougars, Barash 1973b, 1975; black bears, Banfield 1974; golden eagles, Olendorff 1976, Barash 1975, Armitage and Downhower 1974; bald eagles, Beebe 1974; r e d - t a i l e d hawks. Bent 1937).. Two important predators of other marmot species, coyotes (Canis latrans) and g r i z z l y bear fOrsus arctos). are absent from Vancouver island (Cowan and Guiguet 1965) . Whistles were given only 27 of 61 times (56%) that smaller raptors (Faloniformes) and ravens (Corvus corax) were present (Table IV) . whistles also occurred occasionally for other causes, but I could not associate any cause for 59% of the whistles (Table IV) . ;Since such a large proportion of whistling seguences was Table IV. Causes of whistling and keeaw sequences 50 Stimuli Whistling Sequences Number Percent Keeaw Sequences Number Percent Potential Predators black bears 13 6.1 ccugars 3 1.1 la i d eaqles 4 1.9 qclden eagles 5 2.4 unidentified eagles 6 2.8 red-tailed hawks 5 2.4 subtotal 36 Snail Raptors and Havens marsh hawks 10 4. 7 sharp-shinned hawks 4 1. 9 Cooper's hawks 3 1. 4 sparrow hawks 1 0. 5 unidentified small raptors 4 1. 9 ravens 5 2. 4 subtotal Miscellaneous Causes intraspecific chases ai rcraft black-tailed deer people land-tailed pigecns? ccmmon flickers? Canada jays? snow and rock slides sudden h a i l storm subtotal Unknown Causes 27 5 4 4 3 1 2 1 2 1 23 126 2.4 1.9 1.9 1.4 0.5 0.9 0. 5 0. 9 0.5 17.0 12.3 10.8 59.4 14 3 19 2.6 2.6 13.2 13.2 2.6 2.6 5.3 5.3 2.6 36. 8 5. 3 7. 9 50.0 Grand Total 212 100.1 38 100.0 51 not associated with obvious predators, i t i s possible that whistles could have some other function in addition to being alarm c a l l s . Bopp (1955) interpreted the whistles of Harmota marmota as t e r r i t o r i a l c a l l s . However t h i s function has been disputed by Armitage (1962) and Barash (1973b), both of whom emphasized the warning function of th i s c a l l . The reaction of marmots was the same whether or not I was able to associate a cause for them. Therefore, I think that the "unknown" causes of many of the whistles I heard were due to either predators that I did not see or common disturbances such as s l i d i n g snow, rain , wind, h a i l , or moving fog patches that alarmed certain i n d i v i d u a l s (Table IV). I r a r e l y knew which marmot whistled but i n each case when I did the i n d i v i d u a l was already at a burrow entrance. Upon hearing a whistle marmots usually ran to a burrow entrance or to the top of a rock that had a burrow underneath. Marmots rarely ran and entered a burrow d i r e c t l y ; i n f a c t , they frequently emerged from the burrow at the sound of a whistle. Once at a burrow entrance marmots looked about for the cause of the disturbance. Marmots do not immediately enter t h e i r burrows when disturbed presumably because i t i s adaptive to keep any predator in sight and avoid, i f possible, the r i s k of having to subsequently emerge from a burrow when the above ground s i t u a t i o n i s unknown. Emerging from a burrow d e f i n i t e l y involves some r i s k because predators may wait at a burrow entrance u n t i l the occupant emerges. , Coyotes (Canis1atrang) have been observed catching Spermophilus columbianus (Don Bowen 52 personal communication) and M • c a licj at a (I. McT. Cowan personal communication), and foxes {Vuljjes vuljaes) have been observed catching S., undulatus {Gordon Haber personal communication) using t h i s technigue.. Spring hares, south African rodents i n the genus Pedetes, are presumably subjected to the same hunting t a c t i c s . They apparently confound awaiting predators by emerging from t h e i r burrows with a great leap {Vaughan 1972). Marmots usually remained at t h e i r burrows only a few minutes a f t e r the predators had disappeared and whistling had stopped. No " a l l c l e a r " c a l l was apparent. Occasionally some marmots did not appear to react to whistles at a l l , or they merely looked around from where they happened to be at the time. The in t e n s i t y {loudness) of the whistle, and not the whistle duration or i n t e r v a l as suggested by Waring (1966), Barash (1973b) , and Gray (1975), appeared to determine what action marmots would take. Although in t e n s i t y was not measured in the f i e l d , i t was obvious from l i s t e n i n g to whistles that i n t e n s i t y varied greatly. More intense whistles resulted i n more marmots running to burrows. 53 Long whistles were more often associated with t e r r e s t r i a l disturbances, and short whistles were more often associated with a e r i a l disturbances. number of whistling seguences disturbance type long short a e r i a l t e r r e s t r i a l 5 14 25 3 xz=16.81 df=1 p<0.001 n=4 7 Two whistling seguences were dropped from the analysis because there were both long and short whistles i n the sequence. However, both instances were consistent with the above results i f only the f i r s t c a l l of each sequence was considered. Four seguences were dropped from the analysis because the c a l l s were of a medium length. I did not record very detailed observations of the way Vancouver Island marmots reacted to alarm c a l l s . Thus I could not detect any differences between responses to long and short whistles. 54 Keeaws: Vancouver Island marmots produced a sound that has not previously been recorded for other marmots. This vocalization was a f a i n t two-syllable c a l l which sounded l i k e a "kee-aw". The fundamental frequency of a keeaw changed in two stages from 1912 ±39 Hz to 1109 ±57 Hz (Fig 14). There are at least two harmonics, both less intense than the fundamental frequency. The mean duration of a keeaw c a l l was 0.29 ±0.016 s. Keeaws were usually given i n a long ser i e s that varied considerably i n length. On two occasions only single keeaws were given but the mean number of keeaws per sequence was 102 ±27. On one occasion about 900 keeaws were given i n 60 minutes by one i n d i v i d u a l . The i n t e r v a l between keeaws averaged 3.8 ±0.15 s. In long c a l l i n g seguences the i n t e r v a l between c a l l s increased toward the end of the sequence. Keeaws were frequently associated with disturbances and therefore also with whistles (Table IV and Fig 15). However, keeaws were usually given a f t e r whistling had stopped and the predators had l e f t the area. Keeaws therefore seemed to represent a state of "uneasiness" or low i n t e n s i t y alarm. Upon hearing keeaw c a l l s some marmots did not appear to react at a l l , but many marmots ran to a rock or burrow and rested there. Marmots gradually resumed t h e i r previous a c t i v i t i e s within a few minutes after the c a l l i n g started, even when keeaws continued. In a l l but one instance only one animal 55 Figure 14. Representative sonograms of keeaws Recording speed: 19.05 cm per s Playback speed: 19.05 cm per s narrow bandwidth f i l t e r 5000.^ 4500. 4000. 3500.. 3000.. 2500. 5000.J 1500. 1000. 500. 0-0 0*1 0-3 0-5 C7 0-9 TIME (SECONDS) 5000. 4500.. 4000.. 3500.. 3000.. E500.. 5000. 1500-1000. 500-0* 0*0 0.1 0«3 0.5 ~t r 0.7 0>9 1 TIME (SEDLTCE) 56 Figure 15. Frequency with which whistles and keeaws occurred together and separately with and without a known disturbance as a stimulus Light numbers indicate the number of observations dark numbers indicate the percent of the t o t a l vocalization sequences 56a 57 gave keeaws at any one time. Rapid Chirps On one occasion when I approached a marmot colony I heard a rapid series of egually spaced, very short whistles or chirps. I did not record t h i s c a l l , nor did I ever hear i t again. Hisses Individuals in l i v e traps occasionally "hissed" when I approached the trap. When a hiss was given the marmot faced me with i t s mouth open, crouched, and sometimes lunged i n an attempt to bite.„ The hiss spans a wide range of freguencies between 70 and 3000 Hz (Fig 16). }Harmonic structure, i f any, i s very weak. Tooth Chatters A tooth chatter denotes threat i n many rodents {Balph and Balph 1966; Waring 1966,1970; Ewer 1968; Barash 1973b; Brooks and Banks 1973). I heard a Vancouver Island marmot tooth chatter on only one occasion. This occurred when I was handling the animal, an adult male, for tagging. I did not hear either a hiss or a tooth chatter from free ranging marmots. 58 Figure 1,6. Representative sonogram of a hiss Recording speed: 19.05 cm per s Playback speed: 19.05 cm per s Harrow bandwidth f i l t e r F R E O J E N C Y (HZ) 59 Screams and Growls "Screams" and "growls" were occasionally heard during play-f i g h t s and chases but were not recorded, A growl i s a b r i e f low freguency sound probably with a wide range of frequencies. Screams sounded l i k e long high freguency growls. SOCIAL BEHAVIOUR Social Behaviour Patterns I recognized 13 s o c i a l behaviour patterns i n Vancouver Island marmots. Greeting, anal s n i f f i n g , and play-fighting were considered to be r e c i p r o c a l acts because both interactants behaved i n a similar manner when performing these behaviours. Chasing, mounting, a l e r t , avoidance, t a i l r a i s i n g , lunging, suckling, s o c i a l grooming, pl a y - f i g h t i n v i t a t i o n , and play-chasing were considered to be non-reciprocal acts because the behaviour only describes the action of one of the interactants. The description of scent marking i s included in t h i s section both for convienience and because of i t s s o c i a l s i g n i f i c a n c e . G££§£i22 ( G) • Greeting consists of two or more marmots touching t h e i r noses together, or one animal s n i f f i n g the cheek, ear, or (rarely) side of another i n d i v i d u a l . This behaviour pattern has been described for at lea s t f i v e other species of fiariota (M. f l a v i j r e n t r i s , Armitage 1962; M. monax. Bronson 1964; M. oly.mp.us, Barash 1973b; M. c a l i g a t a , Barash 1974b; M. marmota. 60 Barash 1976b) and was the most common s o c i a l behaviour pattern that I observed (Table VI). AfiSi s n i f f i n g (AS). Anal s n i f f i n g consists of two animals standing together with t h e i r bodies p a r a l l e l while nuzzling the anal region of the other marmot. Mounting (Mo). Mounting involved one animal straddling the other from behind with i t s forelegs and placing i t s ventral surface i n contact with the dorsal surface of another. I never observed the dorsal animal thrusting or b i t i n g the back of the other marmot as was seen i n the sexual behaviour of M. f l a v i v e n t r i s (Armitage 1965) and M. olyjngus (Barash 1973b). Suckling (S) and Soc i a l Grooming (SG). Both of these acts were observed occurring only between adult females and t h e i r i nfants. .Avoidance (Av). I recorded avoidance behaviour only when I was sure that an animal's departure was i n response to another i n d i v i d u a l . This occurred when a marmot either ran away during an i n t e r a c t i o n or moved away from an approaching marmot once the l a t t e r was within 3 m. Avoidance does not include animals that were fleeing during a chase. Alert (Al). Alert behaviour was performed by marmots that appeared to be uneasy about the approach of another i n d i v i d u a l . The alerted animal watched the approaching marmot from a r i g i d 61 crouched stance. £1131113 (*•) • Lunging consists of thrusting the front paws foreward toward another marmot, occasionally making contact. l a i i Raising (TR). T a i l r a i s i n g i s a graded display which consists of a marmot erecting the hair on the t a i l and r a i s i n g the t a i l up, noticeably arched. The t a i l may be raised further u n t i l the f l u f f e d t a i l l i e s f l a t along the back of the animal. When t h i s display i s given the t a i l i s usually moved very slowly or held motionless at any point within t h i s range of positions. A l l t a i l r a i s i n g displays were performed at the same time as one of four behaviour patterns: either greeting, anal s n i f f i n g , a l e r t , or play-fighting. However, not a l l of the occurrences of these behaviours were accompanied by t a i l r a i s i n g . Thus, one could consider t a i l r a i s i n g to be an optional component of each of these behaviours rather than as a separate behaviour pattern. Very similar t a i l r a i s i n g behaviour was also observed by Barash (1973b:184,198, and Fig 23) in M. oly.S£H§» by Gray (1967:44 and 50) in M. c a l i g a t a , and by Armitage (1962:325 and Fig 5) and Waring (1966: 181) i n M. f l a v i v e n t r i s . This behaviour i s very d i f f e r e n t from the the rapid pumping and swirling that i s c h a r a c t e r i s t i c of t a i l movements i n M. marmota (Koeing 1957) and from the " t a i l flagging" of H. f l a v i y e n t r i s i n which the arched t a i l i s raised up and waved from side to side (Armitage 1974:243). T a i l r a i s i n g i n H. Sonax also involves rapid t a i l movement (Bronson 1964:471). 62 Chasing (C) and Plav.-cjiasing (PC). Chasing was an agonistic encounter that d i f f e r e d from what I termed play-chasing i n the following ways: 1. Play chases were slower and shorter, usually less than 7-10 metres. 2. Play-chases were always s i l e n t whereas agonistic chases were occasionally accompanied by whistles, sgueals, or growls. 3. Play-chases always ended with the interactants r e s t i n g or feeding close together or i n t e r a c t i n g i n a non-agonistic way. Agonistic chases were ra r e l y followed by other acts since the interactants were usually separate after the chase (Fig 17). Chasing and play-chasing always caused the other marmot to f l e e , thus i t was not necessary to distinguish " f l e e i n g " as a separate behaviour pattern. Play-fighting (PF). Play-fighting consists of two marmots r i s i n g up on th e i r hind legs and pushing with t h e i r forelimbs against the other marmot's chest or shoulders. This behaviour has also been described for M. olympus (Barash 1973b), H. f l a y i v e n t r i s (Armitage 1973, 1974, Barash 1973a), M. caljgata (Barash 1974b), and M. marmota (Barash 1976b). P i a i - f i g j l t I n v i t a t i o n (PI) . A play-fight i n v i t a t i o n was a v i s u a l s i g n a l that appeared to indicate that the marmot performing t h i s behaviour was prepared to play-fight. This posture ranged from simply r a i s i n g the shoulders to r a i s i n g the whole body into a v e r t i c a l p o sition on the back legs. 63 Scent Harking (SH) . , Vancouver I s l a n d marmots f r e q u e n t l y made long sweeping motions with t h e i r cheeks a g a i n s t rocks t h a t were at the entrances to burrows. T h i s behaviour presumably d e p o s i t s scent i n the form of s e c r e t i o n s of t h e i r f a c e glands. Marmots sometimes scent marked a f t e r s o c i a l i n t e r a c t i o n , but scent marking was u s u a l l y performed as an i n d i v i d u a l behaviour. A l l North American and some, i f not a l l , E u r a s i a n marmot s p e c i e s have f a c e glands (Rausch and Rausch 1971:90) and use them f o r s c e n t , marking (M. c a l i g a t a . Gray 1967:48; M. marmota, Koeing 1957:519; Asian marmots, Bibikov 1967 c i t e d i n Rausch and Rausch 1971:90; M. monax and M. b r o w e r i R a u s c h and Rausch 1971:90-91; U. olvmp.us, Barash 1973b: 184; H. f l a v j , v e n t r i s , Armitage 1976b). Dominance R e l a t i o n s h i p s Three behaviour p a t t e r n s , c h a s i n g , avoidance, and l u n g i n g , c l e a r l y i n d i c a t e d the dominant marmot of an i n t e r a c t i o n . The dominant marmot i n a chase was the animal doing the c h a s i n g . Lunging was a l s o c h a r a c t e r i s t i c o f dominant animals s i n c e i t u s u a l l y caused other marmots t o move away. F l i g h t and avoidance t h e r e f o r e , c h a r a c t e r i z e d the subordinate i n d i v i d u a l s of an i n t e r a c t i o n . A dominance matrix based on c h a s i n g , l u n g i n g , and avoidance was c o n s t r u c t e d as d e s c r i b e d by Brown (1975) and i s presented i n T a b l e Va. I t i s apparent from these data t h a t a dominance h i e r a r c h y e x i s t e d i n the form, a d u l t males > a d u l t females > two-year-old females > y e a r l i n g males > y e a r l i n g females. 64 Table V. Dominance matrices of age and sex classes of Vancouver Island marmots (a) . Dominance Matrix Based on the Freguency of Occurrence of Chases, avoidance, and Lunges Between Age:sex Classes Dominant Subordinate Row Column AM* AF 2F 2? YM YF Y? Totals Totals AM 3 2 2 7 1 AF 1 2 7 2 10 2 24 4 :""':"\;.'?2F' 1 1 2 4 7 ' V ''•'i.-.:":''2?'; 0 2 ' . . YM 1 1 2 1 ''l'''T''~i!':"Yt: 0 14 Y? 0 6 (b) . Dominance Matrix Based on the Freguency of Occurrence of Eight Behaviour Patterns (C, Av, Al, L, PI, TR, M, SG) Between Age:sex Classes Dominant Subordinate Row Column AM AF A? 2F 2? YM YF Y? I? Totals Totals AM 23 1 8 4 2 7 3 48 6 AF 2 2 1 13 2 10 3 3 36 29 A? 3 3 3 2F 3 3 1 2 2 3 14 21 2? 1 1 6 YM 1 3 4 4 YF 0 22 Y? 0 9 I? 0 6 * KEY AS IH TABLE I I I 65 When t h e o t h e r t e n b e h a v i o u r p a t t e r n s w e r e a n a l y z e d w i t h r e s p e c t t o t h e d o m i n a n c e r e l a t i o n s h i p s i n d i c a t e d i n T a b l e V a , i t was a p p a r e n t t h a t t a i l r a i s i n g , p l a y - f i g h t i n v i t a t i o n s , m o u n t i n g , s o c i a l g r o o m i n g , a n d a l e r t , w e r e a l s o s t a t u s i n d i c a t i n g b e h a v i o u r s . D o m i n a n t marmots p e r f o r m e d p l a y - f i g h t i n v i t a t i o n s , s o c i a l g r o o m i n g , a n d m o u n t i n g , and s u b o r d i n a t e m a r m o t s p e r f o r m e d a l e r t a n d t a i l r a i s i n g b e h a v i o u r . T h e d o m i n a n c e m a t r i x b a s e d o n a l l 8 o f t h e s e b e h a v i o u r s i s p r e s e n t e d i n T a b l e V b . O f t h e 11 e x c e p t i o n s { r e v e r s a l s ) , f i v e { p o s s i b l y s i x ) o f t h e s e o c c u r r e d b e t w e e n a g e a n d s e x c l a s s e s t h a t were a d j a c e n t i n t h e d o m i n a n c e h i e r a r c h y , where v a r i a t i o n w o u l d be most l i k e l y t o o c c u r . Ho d o m i n a n c e r e l a t i o n s h i p s were a p p a r e n t among t h e o t h e r 5 b e h a v i o u r p a t t e r n s . , T h e r e were no s i g n i f i c a n t d i f f e r e n c e s b e t w e e n t h e number o f d o m i n a n t and s u b o r d i n a t e marmots t h a t i n i t i a t e d g r e e t i n g s { X 2=0 . 7 8 , n=33) , p l a y - f i g h t s (X 2=0.06, n = 1 7 ) , o r a n a l s n i f f s {X 2=2.00, n = 1 8 ) . S u b o r d i n a t e s d i d n o t t e r m i n a t e a n y more g r e e t i n g s (X 2 =1.00, n=25) o r p l a y - f i g h t s (X 2=0.60, n~15) t h a n d i d d o m i n a n t m a r m o t s . 66 The Frequency of S o c i a l Behaviour P a t t e r n s I observed a t o t a l o f 785 b e h a v i o u r a l a c t s o c c u r r i n g i n 587 s o c i a l i n t e r a c t i o n s over the two summers that I observed Vancouver I s l a n d marmots. The freguency t h a t each act was observed between age-sex c l a s s e s i s presented i n Table VI and the r e l a t i v e f r e g u e n c i e s t h a t each behaviour occurred w i t h i n age-sex c l a s s e s i s presented i n Table VII. The data i n Table VI do not i n d i c a t e the a c t o r or r e c i p i e n t i n n o n - r e c i p r o c a l i n t e r a c t i o n s , the t a b l e j u s t shows how f r e g u e n t l y each dyad was observed i n a s p e c i f i c s o c i a l behaviour p a t t e r n (see s e c t i o n on Dominance R e l a t i o n s h i p s ) . The most s t r i k i n g f e a t u r e of the data i n Table VII i s that a l l age-sex c l a s s e s used the same behaviour p a t t e r n s i n approximately the same p r o p o r t i o n s . G r e e t i n g and p l a y - f i g h t i n g were the most common s o c i a l behaviour p a t t e r n s used by Vancouver I s l a n d marmots. .. They accounted f o r 65% of a l l behaviour p a t t e r n s t h a t I observed over the whole study and were c h a r a c t e r i s t i c of the behaviour o f a l l s i x age-sex c l a s s e s t h a t I r e c o g n i z e d . T a i l r a i s i n g was u s u a l l y the next most commonly used behaviour p a t t e r n . The freguency of the other 10 a c t s v a r i e d among d i f f e r e n t age-sex c l a s s e s but they were a l l r e l a t i v e l y r a r e . The degree of s i m i l a r i t y among age-sex c l a s s e s (Table VII) can not be compared s t a t i s t i c a l l y because the data are not independent i . e . g r e e t i n g s o c c u r r i n g between a d u l t males and a d u l t females i n c r e a s e the g r e e t i n g freguency of both of these groups. <0 T a b l e V I . The f r e q u e n c y c f o c c u r r e n c e o f e a c h s o c i a l b e h a v i o u r p a t t e r n between age-sex c l a s s e s DYAE G AS C AV A l TR PF PC PI L Mo SG S TOTAL *(!:AF 33 4 2 2 1 18 16 2 78 A M : 2 F 7 1 3 5 8 1 1 26 AB: YF 6 2 2 ii 9 1 24 AR: YM 4 2 5 11 AM:? 21 5 6 5 2 9 7 1 1 57 AF: AF 2 2 A F: 2 F 41 3 4 1 6 11 3 69 AF: YF 7 2 6 1 5 3 24 AF: YM 8 1 4 13 AF:I 11 3 5 19 AF:? 18 a 9 a 2 7 3 1 2 2 52 2 F: YF 6 1 1 1 1 10 2F: YM 3 1 1 n 9 2F:? 8 2 1 u 15 YF: YF 1 1 n 6 YF: YM 1 1 1 3 Y I : ? 4 1 1 1 7 YM:? 1 1 1:1 16 1 13 2 32 I:? 2a . 1 1 3 4 2 35 ?: ? 70 6 28 6 31 118 30 1 2 292 TOTAL 290 27 60 28 10 88 217 36 7 7 7 3 5 785 M = B a l e 2 = Twc-year-o l d C = C h a s i n g PC = P l a y - c h a s i n g f = Female Y = Y e a r l i n q Av = A v o i d a n c e L = L u n g i n g ? = Sex and I = I n f a n t PF = P.lay- f i q h t i n q Bo = Mo u n t i n g age unknown G = G r e e t i n g ' TR = T a i l R a i s i n g SC = S o c i a l Groominq A = A d u l t S = S u c k l i n g A l = A l e r t AS = A n a l S n i f f i n q PI = P l a y - f i g h t I n v i t a t i o n OO ID T a b l e V I I . P e r c e n t a g e s o f s o c i a l b e h a v i o u r p a t t e r n s per ane-sex c l a s s AGI-SEX BOW TOTAL c i a s s G AS c Av A l TR PF PC P I L Mo SG S TOTAL ACTS AM 36 6 5 4 3 19 23 2 1 2 101 196 AF 46 4 . 10 4 2 12 1 5 <1 1 2 2 1 2 101 259 2F 50 2 2 6 4 10 22 3 1 100 129 YF 33 6 11 4 9 30 1 3 4 101 80 YM 46 3 5 3 5 35 3 100 37 I 57 3 1 3 25 5 3 4 101 118 •» 29 3 10 3 1 11 34 9 <1 1 1 102 751 AVERAGE 37 3 8 4 1 1 1 28 5 1 1 1 <1 1 101 1570 M = Male 2 = Two - y e a r - o l d C = C h a s i n g PC = P l a y - c h a s i n g -F = Female Y = Y e a r l i n g Av = Av o i d a n c e L = Lu n g i n g ? = Sex and I = I n f a n t PF = F l a y - f i g h t i n g Mo = Mounting age unknown G = G r e e t i n g TR = T a i l R a i s i n g SG = S o c i a l Grooming A = A d u l t S = S u c k l i n g A l = A l e r t AS = Anal S n i f f i n g EI = P l a y - f i g h t I n v i t a t i o n 69 The s i m i l a r i t y i n the behaviour of age-sex classes on the whole does not reveal anything about the nature of interactions occurring between s p e c i f i c dyads. I t was not possible to compare the absolute frequencies with which di f f e r e n t behaviour patterns occurred between dyads because the observation times di f f e r e d amonq ind i v i d u a l s (see section on Hates of Social Behaviour). However, what can be compared are the frequencies of occurrence of any given act as a proportion of the t o t a l number of acts. In order to test for s i m i l a r i t y between dyads I compared the r e l a t i v e frequency with which acts occurred between dif f e r e n t dyads (Table VI) using a Chi-sguare test for independence. When i t was necessary to lump the frequencies of certain behaviours to avoid expected values le s s than one, I lumped: 1) chasing, a l e r t , avoidance, and lunging, because of the agonistic nature of these acts (see also Armitage 1962, 1973, 1976a), and 2) greeting and anal s n i f f i n g , because of the cohesive nature of these acts. I f the t o t a l number of acts was less than twenty I used the Fisher Exact Probability test. Thirteen of 15 comparisons were not s i g n i f i c a n t (Table VIII), thus indicating that the behaviour patterns were independent of the interactants. That i s , the behaviour patterns used i n infant : infant interactions were not d i f f e r e n t from the behaviours used between infants and non-infants nor were the behaviour patterns used i n adult male : yearling male interactions d i f f e r e n t from those used i n adult male : adult 70 Table V I I I . Comparisons of the r e l a t i v e freguency with which s o c i a l behaviour p a t t e r n s occurred between d i f f e r e n t age-sex c l a s s e s COMPREISON n X 2 df P AM: AF AM: 2F 104 2.76 4 .60 AM: AF - AM: YF 10 2 3.75 4 .44 AM: YF - AM: YM 35 .49 2 .78 AM: Y F - AM: 2F 50 .35 4 .98 AM: YF - AF: YF 48 11.55 4 .02* AM: YM -• AF: YM 24 1.62 2 .45 AM: 2F - AF: 2F 95 5.49 3 .14 AF: 2F - AF: YF 93 25.96 3 <.01* AF: YF - AF: YM 37 4.6 8 2 .09 AF:2F - AF: YM 82 .17 3 .91 AF: YF - YF: YF 30 4.80 2 .09 2F:YF - YF: YF 16* .96 2F: YM - 2F: YF 19» .42 YF: YF - YF: YM 9* .21 1:1 - I:NI 67 2.59 2 .27 * = P<0.05 1 = F i s h e r Exact P r o b a b i l i t y T e s t , N<20 M = Male 2 = Two-year-old F = Female I = I n f a n t A = Adult NI = Non-infant 71 female or adult male : yearling female interactions. Chi-sguare for the comparisons between adult females and yearling females and other dyads were s i g n i f i c a n t at the 5% l e v e l {* in Table VIII, AF:2F and AH: YF) or the 10% l e v e l (YF: YF and AF: YM, Table VIII). There was more aggression between adult females and yearling females than there was between other dyads. For example, greetings made up a much higher proportion of the interactions between adult females and two-year-old females than between adult females and yearling females (59% vs 29%), while the opposite trend was evident with chases (4% vs 25%). A few dyads were s u f f i c i e n t l y d i f f e r e n t from the general pattern of behaviour that I observed between age-sex classes as a whole, that s t a t i s t i c a l tests were not reguired. Adult males were never seen to interact with other adult males even though there were two i n d i v i d u a l s present on the Haley Lake study area throughout the whole of 1974. Adult females interacted on only two occasions, both of which were chases. Infants experienced s i g n i f i c a n t l y l e s s agonistic behaviour (C, A l , Av, L) than did non-infants (X2=13.59, p<0.0l, df=1) . 72 Interaction Sequences Most of the 587 s o c i a l i n t e r a c t i o n s that I observed during t h i s study consisted of only one behaviour pattern. However, 21% (124 of 587) of a l l interactions consisted of a sequence of two or more acts. The average length of an i n t e r a c t i o n sequence was 2,6 acts (SE=0.J2, range= 2-7). There were no s i q n i f i c a n t differences amonq age-sex classes or s p e c i f i c dyads i n the average number of acts per inte r a c t i o n . Greeting was usually the i n i t i a l behaviour of an inte r a c t i o n sequence. Greetings began 53% of a l l inte r a c t i o n seguences, play-fighting began 30%, and seven other acts i n i t i a t e d the remaining 17% of the seguences. The r e l a t i v e freguencies with which two-act seguences occurred i s i l l u s t r a t e d i n Figure 17. , Since t a i l r a i s i n g always occurred at the same time as some other act, rather than before or af t e r i t , i t was omitted from calculations r e l a t i n g to Figure 17. Host interactions proceeded from a greeting to a play-f i g h t . Some behaviour patterns occurred in interaction seguences much more often than they occurred as single acts. , Twenty-five of 27 anal s n i f f s , 26 of 36 play-chases, and a l l seven play-f i g h t intentions occurred in sequences. 73 Figure 17. Temporal context with which two-act seguences occurred The width of the l i n e i s proportional to the the freguency with which the behaviour sequence occurred The narrowest l i n e represents 2% of a i l seguences 26% of a l l sequences proceeded from a greeting to a play-fight 74 R a t e s o f S o c i a l B e h a v i o u r F i e l d o b s e r v a t i o n s o f b e h a v i o u r r a r e l y a l l o w t h e i n v e s t i g a t o r t o o b s e r v e e a c h a n i m a l o r e a c h g r o u p o f a n i m a l s f o r a n e g u a l amount o f t i m e . T h e r a t e t h a t a b e h a v i o u r p a t t e r n i s p e r f o r m e d i s t h e r e f o r e a more u s e f u l p a r a m e t e r f o r d e s c r i b i n g b e h a v i o u r t h a n i s f r e q u e n c y . R a t e s c a n b e u s e d t o c o m p a r e t h e a b s o l u t e d i f f e r e n c e s i n t h e b e h a v i o u r b e t w e e n i n d i v i d u a l s a n d s p e c i e s , w h e r e a s c o u n t s c a n o n l y b e u s e d t o c o m p a r e t h e r e l a t i v e f r e g u e n c y w i t h w h i c h b e h a v i o u r p a t t e r n s o c c u r r e d w i t h i n t h e b e h a v i o u r o f i n d i v i d u a l s . T h e r a t e s o f a l l b e h a v i o u r p a t t e r n s v a r i e d somewhat b e t w e e n c o l o n i e s o n e a n d two ( F i g 1 8 ) . A l l marmots i n c o l o n y two h a d h i g h e r g r e e t i n g r a t e s t h a n d i d t h e c o r r e s p o n d i n g a g e - s e x c l a s s i n c o l o n y o n e . I a v e r a g e d t h e r e s u l t s o f t h e t w o c o l o n i e s f o r e a c h b e h a v i o u r o f e a c h a g e - s e x c l a s s , t o o b t a i n t h e f i n a l e s t i m a t e o f i n t e r a c t i o n r a t e s o f V a n c o u v e r I s l a n d m a r m o t s . T h e r e was more v a r i a t i o n i n t h e r a t e s o f b e h a v i o u r p a t t e r n s among a g e z s e x c l a s s e s ( T a b l e IX) t h a n t h e r e was i n f r e g u e n c i e s ( T a b l e V I ) . R a t e s o f g r e e t i n g a n d p l a y - f i g h t i n g were h i g h i n a l l a g e - s e x c l a s s , w i t h a d u l t a n d t w o - y e a r - o l d f e m a l e s h a v i n g t h e h i g h e s t g r e e t i n g r a t e s . Y e a r l i n g s h a d t h e h i g h e s t p l a y -f i g h t i n g r a t e s a n d y e a r l i n g f e m a l e s were most o f t e n i n v o l v e d i n c h a s e s a n d l u n g e s . ^The r a t e s o f b e h a v i o u r p a t t e r n s p e r d y a d a r e p r e s e n t e d i n T a b l e X . Two v a l u e s a r e p r e s e n t e d where a p p r o p r i a t e , f o r n o n -75 Figure 18. A comparison of interaction rates between colonies one and two 06 I c in c o 2 . 0 r •1.5 y 1.0 u (TJ (D £ 0 . 5 H colony one • colony two AM AF 2F YM AM AF 2F YM AM AF 2F YM AM AF 2F YM Greeting Play Agonistic Tail Raising Fighting (c, A V , A I , u 76 Table IX. Interaction rate per age-sex class cf behavioural acts per thousand hours Age:Sex Behaviour Pattern :lass G* AS c Av Al TE PF P I L Ho AH 488 513 0 99 160 30 1 138 74 0 24 AF 1225 32 56 32 0 80 265 8 32 74 2F 1005 2 3 22 45 203 44C 271 102 0 79 I F 388 0 388 97 0 0 583 0 291 0 Yfl 606 0 0 0 0 101 303 0 0 0 * KEY AS IN TABLF VI 77 X. Interaction rate per dyad per behaviour per thousand hours DYAD G* AS C Av A l TR PF PI L AM: AF 95 12 2 4 1 49 28 AM:2F 78 4 70 76 26 AM: YF 32 12 10 20 42 A B : Y M 38 35 30 AF: AM - - 4 2 AF: AP 14 A F : 2 F 258 7 12 37 27 AF: YF 30 9 41 6 30 A F : YM 228 61 2F: AM - -2F.-AF - - 3 9 _ 2 F : YF 58 8 11 15 9 2 F : YM 76 15 62 YF: YF 20 73 YF: YM 8 Y M : A F - - 14 _ YM: YF - - 2 7 38 * KFY AS IN TABLE VI 78 'reci.pr.pcal behaviour patterns, but as was noted in the discussion of Dominance Relationships, there were only a few rreversals i . e . yearling males chasing adult females. Rates were highest for greeting and play-fighting i n most dyads. The highest greeting rates occurred between adult and two-year-old females and adult females and yearling males. The dyad with the highest rate of agonistic behaviour was adult to yearling females. For most dyads (9 of 12) the greeting rates exceeded the rates of a l l agonistic acts combined (Fig 19). The rates of a l l behaviour patterns were highest in June and generally declined through July, August, and September (Fig 20). Greeting rates varied less than other behaviour patterns while the rate of agonistic acts varied the most, being much higher i n June than i n a l l other months. Greeting rate was r e l a t i v e l y constant throughout the day, with increases just after emergence from the burrow i n the morning, and just before marmots entered the burrow for the night (Fig 21) . 79 Figure 19. Greeting and agonistic interaction rates per dyad 79a INTERACTION RATE PER THDLJSANO ANlMAL-t-OJRS 6 8 » 6 - S 8 S 8 8 § g S ' B S B S H 1 1 1 1 h H 1 1 1 1 1 1 1 1 > > -n I 80 F i g u r e 20. V a r i a t i o n i n i n t e r a c t i o n r a t e s among months IKTERACTIGN RATE PER ANIMAL-HOUR b o o o o o o o o o K K H - K K ^ H K K K n j f U r u 81 Figure 2 1 . Variation in the greeting rate throughout the day 81a ru GREETINGS PER ANIMAL-HDUR i o o o o o H K K ^ F - r u r u r u -P h P P p — P £ P — p — P P f 82 Dispersion, T e r r i t o r i a l i t y , and Scent Marking On colony two, a l l four marmots had the same home range. The size of t h i s area was approximately 3 Ha. On colony one, the dispersion of marmots was more complex. In May 1974 there was a large amount of home range overlap among a l l animals. Most of t h i s overlap occurred i n the area with steep c l i f f s (Fig 22). U n t i l mid-June t h i s was the only area free of snow, thus i t was the only area where food was available. By the middle of June, most of the snow had melted from the colony and marmots fed on plants that were growing on the c l i f f s much less freguently. At t h i s time the amount of overlap between the home ranges of the two adult females gradually decreased. By July t h e i r ranges were completely separate and they remained that way for the duration of the summer., Adult female #15 occupied the lower half of the colony and adult female #12 occupied the upper h a l f (Fig 23). One adult male, #4, emigrated from colony one to colony two in early June and the remaining adult male, #13, moved over the whole colony, an area of about 4.5 Ha (Fig 22). However, a third, adult male, #17, immigrated to colony one at the end of June and occupied a s i m i l a r area to adult female #12 on the upper 2.2 Ha of the colony. Shortly a f t e r the a r r i v a l of adult male #17, adult male #13 d r a s t i c a l l y reduced his home range u n t i l i t did not overlap at a l l with that of #17 (Fig 24). From July through u n t i l the animals hibernated i n September there 83 Figure 22. Home ranges of the two adult females (#12 and #15) i n May 1974, and the two adult males (#13 and #17) i n June 1974, on the Haley Lake study area The dots i n d i c t a t e the locations of scent markings made by adult male #13 before the a r r i v a l of adult male #17 (n=3). This figure i l l u s t r a t e s the same area shown i n Fig 2B. 83a 84 Figure 23. Home ranges of four female marmots on the Haley Lake study area i n July 1974 Dots indicate the locations of a l l scent marks made by adult female #12 (n=4). C i r c l e s i n d i c a t e the locations of a l l scent marks made by two-year-old female #9 (n=5). Triangles i n d i c a t e the locations of a l l scent marks made by adult female #15 (n=5) . Arrows indicate the locations of chases. #14 i s a yearling female. #17 i s an adult male. 85 Figure 24. Home ranges of the two adult males on the Haley Lake study area in August 1974 Dots indicate the locations of a l l scent marks made by adult male #13 in June, aft e r the a r r i v a l of adult male #17 (n=2). Triangles indicate the locations (n=13) of a l l scent marks (n=17) made by adult male #13 in July. Squares indicate the location of a l l scent marks made by adult male #13 in August (n=1) . C i r c l e s indicate the locations (n=6) of a l l scent marks (n=12) made by adult male #17. 85a 86 were c l e a r l y two main areas on the colony. The upper area was occupied by marmots #12 and #17 and the lower area was occupied by animals #13, #15, two two-year-olds, and three yearlings. Yearling female #14 was the only marmot that consistently used part of both areas (Fig 23). One adult animal occupied peripheral areas and was rarely seen. I analysed the behavioural interactions between occupants of the upper and lower areas of colony one to determine the proximal causes of t h i s pattern of dispersion. Adult females #12 and #15 interacted only on two occasions. Both of these interactions involved #15 chasing #12 from the area normally used only by the occupants of the lower area (Fig 23). Both adult females occasionally scent marked within t h e i r areas of exclusive use (Fig 23). Brown and Orians (1970) accept the concept of a t e r r i t o r y being a defended area and e x p l i c i t l y define defense as being either 1.. actual defense such as chasing away an intruder, or 2. performing i d e n t i f y i n g acts such as scent markings. , Thus my data suggest that adult Vancouver Island marmot females are t e r r i t o r i a l with respect to other adult females, but that their t e r r i t o r i e s are smaller than th e i r home range. I did not see any interactions at a l l between adult males #17 and #13 in spite of the dramatic change i n the home range of #13 that appeared to be d i r e c t l y related to #17*s a r r i v a l . There could have been some rare and s i g n i f i c a n t i nteractions between these two marmots that I missed seeing but usually these 87 two animals just avoided one another. Avoidance was probably enhanced by the deposition of scent marks. Adult males scent marked much more than adult females. The two-year-old female was the only other age-sex class of marmot observed to scent mark. age-sex class freguency of scent marking 42 9 5 The d i s t r i b u t i o n of #13*s scent marks before and aft e r the a r r i v a l of #17 i s shown in Figures 22 and 24. Host of the scent marking that I observed was done by #13 just a f t e r the a r r i v a l of #17 (Fig 25). Adult male #4, on colony two, also increased h i s rate of scent marking i n July (Fig 25). This increase may also have been due to the presence of the scent of another adult male since #4 had just immigrated to t h i s l o c a l i t y . Even though the ranges of #13 and #17 s t i l l overlapped somewhat i n July they each avoided the area enclosed by each others scent marks. The area enclosed by scent marks was also the area of maximum use of each marmot. , I analyzed the amount of time that marmots spent moving but not feeding. Adult males moved s i g n i f i c a n t l y more than other age-sex classes and adult male #13 moved s i g n i f i c a n t l y more than adult males #4 and #17. The r e l a t i v e l y large amount of time spent moving by adult male #13 occurred as he patrolled his home range. P a t r o l l i n g involved moving around the perimeter of the adult males :adult females two-year-old female 88 Figure 25. Rates of scent marking by adult males #4, #13, and #17 Numbers above bars indicate the number of scent marking bouts observed X indicates no data Rate of Scent Marking (number/hr) o > c C o X O 0) - T " do ro IV) 0) CD CO 89 home range greeting other marmots and occasionally scent marking. Adult male #4 did not patrol his home range as often, probably because he never came into contact with other adult males. I think that adult male #17 patrolled less often than #13 because the physical c h a r a c t e r i s t i c s of the habitat allowed him to see most of h i s home range from the tops of the rocks that he freguently rested on. Moving marmots made themselves more conspicuous than feeding marmots by preceeding more of t h e i r movements with t a i l f l i c k s (67% vs 28%, X2=18.3, p<0.01). These observations indicate that adult Vancouver Island marmot males are also t e r r i t o r i a l with respect to other adult males. They occupied fixed areas of exclusive use that could be considered to be defended by scent marks. Interactions occurred between adult female #12 and adult male #13 before the a r r i v a l of adult male #17. These interactions were s i m i l a r to those occurring between adult female #15 and #13, and between #12 and #17 (X 2 = 2.28, p>.25). Thus adult female #12 did not try to defend the upper of the Haley Lake colony against #13, nor did she react aggressively to #1.7. when he f i r s t arrived. As I mentioned e a r l i e r , yearling female #14 consistently moved between the upper and lower areas. However, t h i s i n d i v i d u a l was treated quite aggressively by both #12 and #17, 6 chases (Fig 23) and 3 lunges out of a t o t a l of 20 interactions. In the lower area none of the 12 interactions between #14 and #13, or #14 and #15, were chases or lunges. Some aggression did occur among marmots i n the lower area but i t 90 did. not appear to be related to s p e c i f i c locations and i t appeared as i f a l l occupants could move about f r e e l y anywhere within t h i s area. 91 DISCUSSION VOCALIZATIONS I n t e r s p e c i f i c Comparisons of Marmot Vocalizations The vocalizations of Vancouver Island marmots are very s i m i l a r to those of M. c a l i g a t a and H. olympus (Table XI). However, there are some important differences. Barash (1973b) never heard whistles from M. olyjijgus that were as long as those commonly given by M. vancouverensis. Rapid chirps were often given by M. olympus (Barash 1973b) but I heard them from H« : vancouverensis only once. The keeaw of M. vancouverensis corresponds in M. c a l i g a t a to a "queeuck" c a l l (Gray 1967) and the "low freguency c a l l " described by Taulman (1975). The medium c a l l of M. olympus (Barash 1973b) although d i f f e r e n t i n sound structure appears to be the homologous c a l l i n t h i s species. In the one Olympic marmot c a l l i n g sequence that I heard, the f i r s t few c a l l s sounded exactly l i k e a keeaw. Subsequent c a l l s were pure toned whistles of a medium lenqth as described by Barash (1973b). The duration and i n t e r v a l of medium M. oly_mj>us c a l l s i s quite s i m i l a r to that of keeaws (Table XI). Keeaw, queeuck, low frequency and medium whistle c a l l s were a l l given i n prolonged sequences and a l l three c a l l s seemed to indicate a state of uneasiness. 92 Table XI. A comparison of vocalizations within the Marmota caligata Group Call Characteristics M. vancouverensis M. caligata M. C l 2 1 £ U § ( S > Call Name Duration (s) Interval (s) Freguency (Hz) long whistle* 0.57 17.9 2910 lonq call*< »'> .56-.75<i23> 13.6-16<»*> 280 0-3200<123) Call flame Duration (s) Interval (s) Freguency (Hz) medium whistle* 0.31 6.5 2910 descending and ascending cal l * * ' ) 0.3{to 0.5)<i> 3<i > 3500< » > lcnq c a l l 0.39 >5 2700 Call Name short whistle* alarm chiip*< 1 3> medium c a l l * Duration (s) 0. 22 0. 1 " ' 0.2 Interval (s) 2.9 1.3<»> 1-3 Freguency (Hz) 2910 2500-3200<»3> 2700 Call Naire rapid chirps accelerating chirp< 13> short c a l l * Duration (s) < « 0. 22 0. 1-var iatle< i > . 0.095 Interval (s) < « 2.9 .05-var iable<»' 0.36 Freguency (Hz) ? 2500-3200<i> 2700 Call Name keeaw* queeuck whistle*•> Duration (s) 0.29 0.3<»> Interval (s) 3.8 1<*> Freguency (Hz) 1900-1100 2000-1500<»> * - c a l l s used in correlation analysis cn paqe 93 <» > Tauluian 1975 <*> Warinq 1966 <3> Pattie 1967 <•> Gray 1967 <s> Barash 1973b 93 The mean i n t e r v a l between whistles of M. vancouverensis. M« c a l i g a t a , and M. olympus follows the same pattern., The c a l l i n t e r v a l was p o s i t i v e l y correlated with the duration of the whistle {r=0.91i* n=9 p<0.01. Table XI). This r e l a t i o n s h i p may res u l t from the ef f e c t of vocalizations on respira t i o n . More time may be reguired to recoup expended oxygen after a long whistle than after a short one. M. f l a y i v e n t r i s vocalizations d i f f e r from those of the M. c a l i g a t a group i n that the c a l l i s very short, 0.037 s and does not vary in duration when the i n t e r v a l changes (Waring 1966). M. f l a v i y e n t r i s gives an accelerando chirp (personal observation) that i s sim i l a r to a c a l l described for J|. c a l i g a t a (Gray 1967). This c a l l was not heard from M. vancouverensis or M. olympus (Barash 1973b). I»loyd (1972) described two vocalizations of H. monax. One i s a "short simple whistle" and the other i s a "two-part whistle consisting of a single high i n t e n s i t y shriek followed by a l e s s intense warble." The simple whistle may be quite s i m i l a r to the whistles given by species in the M. c a l i g a t a group, but no sonogram was presented. The two-part c a l l does not resemble any c a l l s so f a r desrcibed f o r any other marmot species., 94 altruism and Marmot Alarm C a l l i n g a l t r u i s t i c behaviour can be defined as behaviour that benefits another organism, not closely related, while being detrimental to the organism performing the behaviour. Benefit and detriment are defined in terms of the contribution to an animal's f i t n e s s where f i t n e s s i s measured by the proportion of an animal's genes l e f t i n the the population gene pool (Pianka 1974). Since a marmot giving an alarm c a l l may attra c t the attention of a predator and thereby subject i t s e l f to a greater r i s k of predation than i f i t had remained s i l e n t , marmot alarm c a l l i n g appears to be a l t r u i s t i c . However, true altruism i s v i r t u a l l y unknown (Pianka 1974) and i f present would be very d i f f i c u l t to account for by natural s e l e c t i o n . There are three ways to account for the evolution of marmot alarm c a l l i n g . The f i r s t explanation i s kin select i o n . Kin selection i s the evolution of c h a r a c t e r i s t i c s within an i n d i v i d u a l that favour the survival of i t s close r e l a t i v e s but that are not necessarily b e n e f i c i a l to that i n d i v i d u a l . Kin selection could account for the evolution of alarm c a l l i n g even i f i t involved some r i s k to the c a l l e r and i f i t i n c i d e n t a l l y benefitted some unrelated i n d i v i d u a l s . Kin se l e c t i o n has been considered an important sel e c t i v e force i n the evolution of alarm c a l l i n g i n birds (Hamilton 1964, Maynard Smith 1965, Emlen 1973). The s o c i a l organization of Vancouver Island marmots i s such that the in d i v i d u a l s receiving the alarm are l i k e l y to be cl o s e l y related to the c a l l e r . Thus, kin selection could 95 account for the existence of alarm c a l l i n g in Vancouver Island marmots. Secondly, alarm c a l l i n g may be selected for because there i s a dire c t benefit to the c a l l e r associated with c a l l i n g . The in d i v i d u a l could benefit from c a l l i n g i f i t confused the predator (Maynard Smith 1965) or i f i t "manipulated" other marmots so as to make the c a l l e r less vulnerable than the other individuals (Charnov and Krebs 1975). Marmot c a l l s could be manipulating other marmots by stimulating them to run to a burrow. The reacting marmots thus become more conspicuous (to me and presumably to predators as well) than the c a l l e r , who has already moved to a burrow entrance. As Charnov and Krebs (1975) argue, the ind i v i d u a l s reacting to the c a l l "use the information to t h e i r own benefit, but by doing so make i t possible f o r the c a l l e r to benefit even more." Trivers (1971) presented other arguments which would account for the evolution of alarm c a l l i n g by di r e c t selection for the i n d i v i d u a l c a l l e r . , He argues that i t i s disadvantageous for an i n d i v i d u a l to have a predator k i l l a nearby conspecific because the predator may then be more l i k e l y to k i l l him i n the near future. This could occur i f the predator was more l i k e l y to (1) develop a search image (Emlen 1973) for that prey species, (2) learn the habits of the prey species and perfect i t s hunting techniques on i t , and 96 , (3) frequent the habitat of the prey species. Giving alarm c a l l s thus tends to prevent a predator from s p e c i a l i z i n g on the c a l l e r ' s species and l o c a l i t y , thereby favouring the i n d i v i d u a l c a l l e r even though c a l l e r s are i n c i d e n t a l l y a l t r u i s t i c to t h e i r non-calling neighbours (Trivers 1971, Charnov and Krebs 1975). There may be another direct advantage to marmot alarm c a l l i n g . I t may be advantageous for an animal to have conspecifics around regardless of how closely related they are. The presence of conspecifics may be advantageous with respect to finding a mate or increasing winter s u r v i v a l due to the reduction of heat loss when animals hibernate i n a group. Thus, there would be selection for alarm c a l l i n g i f the benefit derived from having other animals around exceeds the cost of c a l l i n g . Lastly, alarm c a l l i n g could be brought about by group sel e c t i o n , which i s selection favouring the su r v i v a l of the group as a whole rather than the i n d i v i d u a l (Wynne-Edwards 1962). Group selection could account for a t r u l y a l t r u i s t i c t r a i t to evolve; however, i f group selection exists at a l l i t i s rare (Lewontin 1970), less e f f i c i e n t than i n d i v i d u a l selection (Sterns 1976)i and i t should not be invoked i f a simpler solution (direct or kin selection in the case of marmot alarm ca l l i n g ) exists (Hilliams 1966) . The above arguments were presented to account for the 97 apparent cost (loss of fitness) associated with alarm c a l l i n g . However, no information e x i s t s for any species that there i s any net cost involved with alarm c a l l i n g (Brown 1975). Observations °f Marmota species do not support the sugestion that the alarm c a l l e r i s incurring any r i s k . Vancouver Island marmots appear to protect themselves before whistling. In a l l cases I observed,; the c a l l e r was already at a burrow entrance. Barash (1975) saw eight instances of predation on various Marmota species. Not one of these occasions was preceded by alarm c a l l s from the victim. From the above arguments I conclude that there i s no true altruism associated with marmot alarm c a l l i n g . , If there i s any cost at a l l associated with c a l l i n g , benefits from dir e c t or kin selection would be strong enough to select for alarm c a l l i n g . The Evolution of Whistle Structure Marler (1955, 1957) was the f i r s t to point out how the physical properties of avian alarm c a l l s could be related to t h e i r function. He observed that some bird species have convergently evolved alarm c a l l s that have c h a r a c t e r i s t i c s that appear to make them d i f f i c u l t to locate. Konishi (1973) tested the l o c a t a b i l i t y properties of d i f f e r e n t sound c h a r a c t e r i s t i c s using, barn owls (3?y_to. alba) , a species that i s adapted for lo c a t i n g i t s prey by sound. His r e s u l t s indicated that barn owls locate sound by comparing the intensity of sound between ears. Binaural differences i n i n t e n s i t y r e s u l t from a shadow 98 being cast by the head. Wide bandwidth noises are easier to locate than pure tones because they consist of many frequencies, each of which can be used for binaural i n t e n s i t y comparisons. L o c a t a b i l i t y by t h i s method i s also d i r e c t l y related to freguency (at least between 3 and 9 kHz)., Thus, barn owls located narrow bandwidth pure tones of 3 kHz les s accurately than any of the other sound c h a r a c t e r i s t i c s tested with bandwidth being the primary c h a r a c t e r i s t i c determining the l o c a t a b i l i t y of a sound. Vancouver Island marmot alarm whistles have the precise c h a r a c t e r i s t i c s of the most d i f f i c u l t - t o - l o c a t e sound tested by Konishi (1973). Narrow bandwidth alarm c a l l s are also found i n many other (but by no means a l l ) species of medium si z e , diurnal mammals (Table XII). A d i f f i c u l t - t o - l o c a t e alarm c a l l would presumably reduce the r i s k of predation to the c a l l e r . Thus predation pressure could se l e c t for the evolution of such alarm c a l l c h a r a t e r i s t i c s i n a l l of the species l i s t e d . The alarm c a l l s of the birds recorded by Marler (1955, 1957) a l l had narrow bandwidths. I t appears therefore that there has been remarkably strong convergence for the same alarm c a l l c h a r a c t e r i s t i c s i n both birds and mammals. Selection for d i f f i c u l t - t o - l o c a t e alarm c a l l s does not indicate anything about the f i t n e s s of in d i v i d u a l s giving d i f f i c u l t or easy to locate c a l l s r e l a t i v e to no c a l l . Thus, selection for s p e c i f i c c a l l c h a r a c t e r i s t i c s does not enter into the discussion of the a l t r u i s t i c nature of marmot alarm c a l l i n g (non-callers vs ca l l e r s ) . 99 The Evolution of Whistle Function M* vancouverensis alarm c a l l s not only indicate that there i s a predator around but also what type of predator i s present. The presence of s i t u a t i o n s p e c i f i c alarm c a l l s has not been shown for any other marmot species., Such alarm c a l l s are r e l a t i v e l y common among ec o l o g i c a l l y related species. Avian and t e r r e s t r i a l predators evoke s p e c i f i c alarm c a l l s i n at least four species of ground s g u i r r e l s (S. beechexi, Owings et a l 1977, Fitch 1948; S. armatus, Balph and Balph 1966; •=>. SSSJSiatus, Melchior 1971; S. b e l d i n g i , Turner 1973), mountain viscachas (Laqjdjum peruanum, Pearson 1948), and perhaps black-tailed p r a i r i e dogs (Cynomys ludovicjanus; King 1955, Waring 1970, Smith et a l 1977). fl. O I Z I J J U S (Barash 1973b), M. f l a v i v e n t r i s (Armitage 1962 and personal communication. Waring 1966), and M. caligata (Gray 1967) do not appear to have s i t u a t i o n s p e c i f i c c a l l s . I f s i t u a t i o n s p e c i f i c alarm c a l l s are communicating s p e c i f i c information then the receivers of t h i s information should exhibit a b i o l o g i c a l l y appropriate response. Of the eight species that have s i t u a t i o n s p e c i f i c alarm c a l l s , only two, Belding's ground s q u i r r e l (Turner 1973) and bl a c k - t a i l e d p r a i r i e doqs (King 1955) were observed to react d i f f e r e n t l y to each c a l l . I think that the lack of observations of s p e c i f i c responces in the other s i x species i s probably due to the d i f f i c u l t y in detecting small differences between responses. For example, H. vancouverensis responds to both long and short 100 whistles by running to a burrow and looking about for the cause of the disturbance. I t would be advantageous in t h i s s i t u a t i o n i f whistle length could provide information on where to begin looking for a predator, either i n the a i r or on the ground. I f th i s was the only variation i n the response,it would be very d i f f i c u l t to detect or to test f o r . A e r i a l predators probably exert the greatest selection for d i f f i c u l t - t o - l o c a t e alarm c a l l s . In a l l f i v e species that have both s i t u a t i o n s p e c i f i c and d i f f i c u l t - t o - l o c a t e alarm c a l l s (Table XII), avian predators evoked alarm c a l l s that were less locatable than the corresponding c a l l evoked by t e r r e s t r i a l predators. Avian predators may s e l e c t more strongly for d i f f i c u l t - t o - l o c a t e alarm c a l l c h a r a c t e r i s t i c s because t h e i r range and speed of attack i s greater than that of t e r r e s t r i a l predators. .The presence or absence of d i f f i c u l t - t o - l o c a t e and s i t u a t i o n s p e c i f i c alarm c a l l s i s probably a function of the predation pressures and the variety of possible escape responses that a prey species possesses., As avian predation pressure increases there i s increased s e l e c t i o n for d i f f i c u l t - t o - l o c a t e alarm c a l l s . Situation s p e c i f i c alarm c a l l s w i l l only be selected for when there i s s e l e c t i v e pressure from both t e r r e s t r i a l and avian predators and i n d i v i d u a l prey can increase t h e i r chances of s u r v i v a l by responding d i f f e r e n t l y to these dif f e r e n t predators. 101 Table XII. A l i s t of mammalian species that have narrow bandwidth alarm c a l l s Species Referenee M* Xancouverensis< 2> M..olympus M. ca l iga ta M. fonax Spermophilus armatus<z > S« £ i cha£dson i i S. f r a n k l i n i i S. nndulatus< 2> feeechey.i<2> rock hyraxes: Hyracoidea i'SflU-iiil peruanum< 2> t a g i d i a i boxi Octodon degus C h i n c h i l l a l an iqer Pagyomys fiilorides Po i i cho t i s pataqonum IJeSiolagus s a l i n i g o l a ? § c t i n a t o r sp. Mjcrocayia spp. Cavia spp. t h i s study Barash 1973b Waring 1966 Lloyd 1-972 <»•> Balph and Balph 1966 Banfield 1974<*> Banfield 1974<»> Melchior 1971 Owings et a l 1977 Mathews 1971<i> Pearson 1948 and Eisenberg 1974 Rowlands 1974<»> Eisenberg 1974< »•> Eisenberg Eisenberg Eisenberg Eisenberg George 1974<»> Rood 197 2: 16<»> Rood 197 2: 17<D 1974< »> 1974< * y 1974< i > 1974< * > <ir - No sonograms were presented but the verbal description appears to be appropriate, and a narrow bandwidth sound i s r e l a t i v e l y easy to describe. (z) - Alarm c a l l s are also predator-specific. 102 In addition to warning other marmots to seek cover, whistles may reduce predation by i n h i b i t i n g attacks. I f the predator i s aware that i t has been detected i t may abandon an otherwise auspicious attack because of the o v e r a l l low p r o b a b i l i t y of capturing alerted prey. Alcock (1975) and Brown (1975) noted that alarm c a l l s or signals may i n h i b i t predator attacks. Warning c a l l s may also function as mobbing c a l l s i n birds by discouraging a predator from remaining i n the v i c i n i t y (Wilson 1975). ,. INTERSPECIFIC COMPARISONS OF MARMOT SOCIAL BEHAVIOOS A Test of Barash*s Hypothesis ; In order to test the data obtained in t h i s study against Barash*s hypothesis of marmot s o c i a l i t y , i t was f i r s t necessary to obtain e x p l i c i t predictions pertaining to Vancouver Island marmots. The parameter necessary to make such predictions i s an estimate of the length of the vegetative growing season, as measured by the number of f r o s t - f r e e days i n the absence of snow cover (Barash 1973b). On the Haley Lake study area the number of f r o s t - f r e e days was 115 days i n 1973 and 135 days i n 1974 (Table I I ) . The average number of f r o s t - f r e e days i n t y p i c a l subalpine parkland habitat i s 114 days (Brooke et a l 1970). The number of f r o s t - f r e e days was equivalent to the vegetative growing season i n 1973 but i n 1974 large amounts of snow persisted for two weeks afte r the l a s t f r o s t . This reduced the vegetative growing season to about 121 days. Snow t y p i c a l l y 103 pe r s i s t s f o r about 3 weeks after the l a s t f r o s t i n subalpine environments {Brooke et a l 1970), thus averaqe growing season i n subalpine environments i s about 93 days. The 93 to 121 day growing season i s r e l a t i v e l y long compared to that experienced by other marmot species. M. olympus experiences a short growing season of 40 to 70 days, H. fla.yj.vejntr.is experiences an intermediate growing season of 70 to 100 days, and H. monax i n central Pennsylvania experiences a very long growing season of about 150 days (Barash 1974a). Barash's hypothesis would therefore predict that the s o c i a l behaviour and s o c i a l organization of H. vancouverensis should be very s i m i l a r to that of M.,flaviventris. That i s , Vancouver Island marmots should 1) be s o c i a l l y i n t o l e r a n t as indexed by having a very low greeting rate, 2) be moderately aggressive, 3) have i n d i v i d u a l t e r r i t o r i e s or r e l a t i v e l y d i s t i n c t home ranges, and 4) grow quickly and disperse at an early age, probably as yearlings. Individuals belonging to a highly integrated and stable s o c i a l group would be l i k e l y to greet more often than in d i v i d u a l s belonging to a less cohesive society because greeting presumably reinforces i n d i v i d u a l recognition and perhaps also functions as a method of scent sharing (King 1955; Barash 1973a, 1973b, 1974b; Steiner 1975). Individual recognition on the basis of scent has been demonstrated in many mammalian species (Halpin 1974, Shorey 1976). Scent sharing has been postulated for marmots, ground s q u i r r e l s (Spermophilus spp.), and p r a i r i e doqs (Cynomys spp., Steiner 1975) and for mountain sheep (Ovis spp., Geist 1971). 104 A comparison of Vancouver Island marmot greeting rates with those pf other marmots reveals that Vancouver Island marmots are among the most s o c i a l of a l l marmot species {Fig 26). The greeting rate of M. vancouverensis i s much higher than a l l three estimates of the rate at which M. f,layjyentris greets. This r e s u l t i s inconsistent with the prediction based on Barash's hypothesis. The greeting rate of M. marmota (Barash 1976b, Tables II S IV) appears to be much higher than that of yellow-b e l l i e d marmots (Barash 1973a, Fig3); however, Barash (1976b) states that the greeting rates of M. maoiot§ are s i g n i f i c a n t l y lower. Thus the rates of both greetings and chases i n M. marm.ota are somewhat suspect, at least the way I am interpreting them. The only behaviour pattern that Barash guantified was chasing (Barash 1973a, 1974b, 1976b) and he did not even present that data for H. olympus (Barash 1973b) . I assumed the same chasing rate for M. olympus as M. marmota since Barash (1976b) states that the freguency of chasing did not d i f f e r s i g n i f i c a n t l y between these two species. Thus the only agonistic behaviour that I could compare among most marmot species was the chasing rate. Species with l e s s s o c i a l tolerance would l i k e l y p a r t i c i p a t e in chases more freguently than s o c i a l l y tolerant species. Chasing rate varied l e s s among marmot species than did greeting rates, with H. vancouverensis having a r e l a t i v e l y low rate of chasing (Fig 27). However, the r a t i o of greetings to chases i s probably the most meaningful 105 Figure 26. & comparison of greeting rates among marmot species 1 _ Marmota vancouverensis (this study) 2 - I~ olympus (Barash 1973b) 3 ~ M. c a l i g a t a (Barash 1976b) 4 - M. marmota (Barash 1974b) 5a - H. f l a v i v e n t r i s (Armitage 1974, 1976a) 5b - M. f l a v i v e n t r i s (high elevation, Barash 1973a) 5c - M. f l a v i v e n t r i s (medium elevation, Barash 1973a) 6 - M. monax (Bronson 1964) O 106 paramater to compare among marmot species. There are three reasons for t h i s . F i r s t , s o c i a l l y intolerant species would be expected to have agonistic acts making up a higher proportion of th e i r s o c i a l interactions. Second, a comparison of the rate r a t i o s should overcome the differences among authors in the methods used to calculate i n t e r a c t i o n rates. Third, a comparison of rate r a t i o s i s independent of any e f f e c t s of colony s i z e . When the rates of greeting to chases were compared, Vancouver Island marmots appeared to be the most s o c i a l marmot species (Fig 28), a r e s u l t inconsistent with Barash's hypothesis., The t h i r d prediction of Barash's hypothesis i s that Vancouver Island marmots should have in d i v i d u a l t e r r i t o r i e s or r e l a t i v e l y d i s t i n c t home ranges. This was c l e a r l y not the case. Vancouver Island marmots have completely overlapping home ranges within a colony or, in the case of colony one, within a section of a colony. Complete home range overlap among colony members was also found i n M. olympus (Barash 1973b) and M. ca l i g a t a (Barash 1974b). Yellow-bellied marmots are grouped into harems but may u t i l i z e i n d i v i d u a l l y d i s t i n c t feeding areas (armitage 1962, 1974), while woodchucks are e s s e n t i a l l y s o l i t a r y (Bronson 1964). , The f i n a l prediction of Barash's hypothesis i s that Vancouver Island marmots should grow and mature guickly, and disperse at an early age, presumably as yearlings. Growth was r e l a t i v e l y slow, as yearlings of both sexes, two-year-old 107 Figure 27. A comparison of chasing rates among marmot species 1 ~ Marmota vancouverensis {this study) 2 ~ !!• olxmjDus {see text) ! 3 - M. c a l i g a t a (Barash 1976b) 4 - M. marmota (Barash 1974b) 5 a ~ M« f l a v i v e n t r i s i Armitage 1974f Barash 1976a) ':; 5b - M. f l a v i v e n t r i s (high elevation, Barash 197 3a) :; 5c - M. f l a v i v e n t r i s (medium elevation, Barash 1973a) ! 6 - M. monax (Bronson 1964) 05 o JZ I c CTJ tf) C O '•+-» U (? (TJ O) c (TJ JZ U Q - 0 . 0 h - 1 . 0 h -2.0 h 108 F i g u r e 28. A comparison of the r a t i o o f g r e e t i n g s to chases among marmot s p e c i e s 1 - Marmota vancouverensis ( t h i s study) 2 - |f. o i l ! p u s (Barash 1973b) 3 - M. c a l i g a t a (Barash 1976b) 4 - a. marmot.a (Barash 1974b) 5a - M. f l a v i v e n t r i s (Armitage 1974, 1976a) 5b - M. f l a v i v e n t r i s - (high e l e v a t i o n , Barash 1973a) 5c - M. f l a v i v e n t r i s (medium e l e v a t i o n , Barash 1973a) 6 - M. monax (Bronson 1964) 108a CD H—' 03 cr U) c ro U -*-» CTj O) c V CD o Q I c CO if) c o CD 1.0 0 . 0 2 -1.0 - 2 0 All Age and Sex Classes Combined 109 females, and perhaps also two-year-old males, were distinguishable from adults by weight (Fig 4). An index of growth rate can be obtained by comparing marmots after emergence from th e i r f i r s t hibernation (yearlings) with the weight of adults at that time of year (Table XIII) . Vancouver Island marmots grew slower than both M. f l a v i v e n t r i s and M. monax (Table XIII). Vancouver Island marmots appear to have delayed maturity equivalent to that of M. olympus. Two Vancouver Island marmots that I was sure had dispersed (males #4 and #17) were two years old or older, while neither of the two two-year-old females on my . study areas produced l i t t e r s . . However, none of the adult females on colonies one or two had l i t t e r s in 1974 either. I f either my presence on the colony, my disturbance through trapping or the late spring i n 1974 were responsible for the adult females not breeding then the same factors would be acting on both two-year-old and adult females. , If any of these three reasons were responsible for adult females not breeding then I would have no basis f o r postulating that two*year-old females do not breed because they are immature. However, these three factors can be discounted: 1) Three females produced l i t t e r s in 1973 when I observed but did not trap. Therefore my presence on the colony was not s u f f i c i e n t to i n h i b i t breeding. 2) I trapped one adult female on the Ski Club colony (colony 5) i n 1974 and she s t i l l produced a l i t t e r , thus trapping does not appear to be s u f f i c i e n t to i n h i b i t breeding. 3) The persistence of snow late into the spring of 1974 was not s u f f i c i e n t to i n h i b i t breeding 110 table XIII. a comparison of the r e l a t i v e growth rates of ? yearling marmots Species Yearling/adult Height in the Spring (%) Reference 18 Barash 1976a 44 t h i s study 30 Barash 1973b 50 J. Donaldson in prep 56 armitage et a l 1976 75 Barash 1973a 70 Barash 1973a 65 Snyder et a l 1961 M... calicjata M; vancouverensis H. olympus -M. f l a v i v e n t r i s M." f l a v i v e n t r i s 1 • f l a v i v e n t r i s 1 H« f l a v i v e n t r i s 2 M. monax i high elevation colony z medium elevation colony 111 since breeding occurred on other colonies (e.g. Ski Club and Buttler Peak) even though the snow conditions were very s i m i l a r i n a l l areas. The most l i k e l y explanation for the f a i l u r e of adult females on colonies one and two to breed i n 1974, i s that Vancouver Island marmot females may only breed in alternate years, a reproductive strategy s i m i l a r to M. olympus. Biennial breeding could account for the absence of l i t t e r s in 1974 because jthree l i t t e r s had been born the previous year. Since marmots were not tagged in 1973, I cannot be sure that a l l three that produced l i t t e r s i n 1973 were the same ones present i n 1974 but one d i s t i n c t i v e l y marked ind i v i d u a l was present i n both years. ,None of the four predictions of Barash*s hypothesis were consistent with the observed data on M. vancouverensis. Vancouver Island marmots in a l l instances most cl o s e l y resembled M. olympus, the species with the shortest growing season. Inconsistent r e s u l t s such as these lead one to either t r y to construct a new hypothesis, having rejected the old one, or to modify one or more of the assumptions of the existing one. The most obvious assumption to reject i s , that vegetative growing season i s a useful parameter for indicating the degree of s o c i a l tolerance a marmot species should exhibit. The fundamental guestion that Barash was try i n g to answer when considering growing season was why some marmot species take longer than others to reach adult s i z e . He found that among M. f l a v i v e n t r i s , M. monax, and M. olympus the length of the growing season correlated with the length of time marmots took 112 to mature. However, the r e s u l t s of t h i s study and those of Anderson et a l (1974) are inconsistent with t h i s trend. Vancouver island marmots do not appear to mature (breed) u n t i l t h e i r fourth summer with a 93-121 day growing season while yellow-bellied marmots mature one year e a r l i e r when the growing season i s only 60-100 days (Barash 1974a, Anderson et a l 1974). Thus when more data are considered, vegetative growing season does not appear to be a meaningful parameter a f f e c t i n g how guickly marmots reach maturity. Anderson et-al-(1974) suggest that the length of time that marmots grow throughout the summer (the marmot growing season), would be a more appropriate measure of environmental severity. However, the length of the marmot growing season does not correlate any better with the time taken to reach maturity than does the vegetative growing season. . . . . . . i , Olympic marmots gain weight for up to 120 days (Barash 1973b) while yellow-bellied marmots at high elevations grow f o r only 96 days (Anderson et al 1974) and mature one year e a r l i e r . Neither the marmot growing season nor the vegetative growing season are correlated with the age of maturity because the time taken to reach adult size i s determined by the combination of three independent factors; 1) the rate that marmots put on weight i n terms of grams per day, 2) the number of days that marmots continue to gain weight (the marmot growing season), and 3) the absolute weight of an adult marmot of the species being considered. These three fac t o r s varied among species but I could not determine any consistent unifying trends among them. The most obvious way to determine how long i t takes a 113 marmot to reach adult size i s to measure the growth rate d i r e c t l y . There i s a s i g n i f i c a n t negative c o r r e l a t i o n between the logarithm of the greeting rate and the weight of immature marmots 1} at the end of t h e i r second summer, expressed as a percent of the f a l l weight of an adult <r=0.86, n=6, p=0.03) and 2) the logarithm of the weight of yearlings a f t e r spring emergence from th e i r f i r s t hibernation r e l a t i v e to the weight of adults at that time of year (r=0.95, n=7, p=0.0 01, and Table XIII).. Thus, by measuring the growth rate d i r e c t l y rather than estimating i t from the length of the vegetative growing season, r e l a t i v e s o c i a l tolerance can be predicted f o r a l l marmot species. The reason that Vancouver Island marmots take a long time to mature i s not that they grow slowly in terms of grams per day or because they only gain weight for a short period each summer, for they are intermediate among marmot species with respect to both of these parameters. Vancouver Island marmots take a long time to mature because they have a r e l a t i v e l y large adult body s i z e . A large body s i z e may be selected for because i t increases the chance of successful dispersal. Vancouver Island marmots l i v e i n islands of subalpine habitat and the probability of successful d i s p e r s a l to new habitats i s probably low. However, the p r o b a b i l i t y of successful dispersal probably increases with body s i z e . This would occur i f larger animals could t r a v e l further and faster, and survive for a longer period of time without food and shelter than could smaller animals. I f aggression from adults causes dispersal {Barash 1974a), then 114 s o c i a l tolerance must increase along with any increase in the optimum body size of emigrants, so that undersized animals are not forced to disperse. 115 LITERATURE CITED Alcock, J. 1975. Animal Behavior: An Evolutionary Approach. Sinauer Associates Inc., Sunderland, Mass. 547 p. Altmann, J. 1974. Observational study of behaviour: Sampling methods. Behaviour 49: 227-267., Anderson, D. C., K. B. Armitage, and R. S. Hoffmann. 1976. Socioecology of marmots: Female reproductive strategies. Ecology 57: 552-560. Armitage, K. B. 1962. So c i a l behaviour of a colony of yellow-b e l l i e d marmots (Marmota f l a v i v e n t r i s ) . Anim. Behav. 13: 319-331. Armitage, K. B. 1965. Vernal behaviour in the yellow-bellied ; ; marmot (Marmota f l a v i v e n t r i s ) . Anim. Behav. 13: 59-68., Armitage, K. B. 1973., Population changes and s o c i a l behavior ".' following colonization by the yellow-bellied marmot. J . Mammal. 54: 842-854. Armitage, K. B. 1974. Male behaviour and t e r r i t o r i a l i t y i n the yellow-bellied marmot. J. ,-Zool. Lond. 17 2: 233-265. Armitage, K. B. 1976a. Social behavior and population dynamics of marmots. Oikos 26: 341-355. Armitage, K. B. 1976b. Scent marking by yellow-bellied marmot. J. Mammal. 57: 583-584. Armitage, K. B., and J . F. Downhower. 1974. Demography of yellow-bellied marmot populations. Ecology 55: 1233-1245. Armitage, K. B., J . F. , Downhower, and G. E. Svendsen. 1976. Seasonal changes in the weights of marmots. Am.;Midi. Nat. 96: 36-51. Balph, D. M., and D. F. Balph. 1966. Sound communication of Uinta ground s g u i r r e l s . J. Mammal. 47: 440-450., 116 Banfield, A. W. F. 1974.,The Mammals of Canada. Oniviversity of Toronto Press, Toronto, Ont.,438 p. Barash, D. P. 1973a. Social variety i n the yellow-bellied marmot (Marmota f l a v i v e n t r i s ) . Anim. Behav. 21: 579-584. Barash, D. P. 1973b. The s o c i a l biology of the Olympic marmot. ' A n i m . Behav. Monogr. 6: 171-245. Barash, D. P. 1974a. The evolution of marmot s o c i e t i e s : A general theory. Science 185: 417-420. Barash, D. P. 1974b. Social behaviour of the hoary marmot (Marmota c a l i g a t a ) . Anim. Behav. 22: 256-261. Barash, D. P. 1975. Marmot alarm c a l l i n g and the guestion of a l t r u i s t i c behavior. Am. Midi. Nat. 94: 468-47 0. Barash, D. P. 1976a. Pre-hibernation behavior of f r e e - l i v i n g hoary marmots, Marmota c a l i g a t a . J . Mammal. 57; 182-185. Barash, D. P. 1976b. Social behaviour and i n d i v i d u a l i t y i n free-l i v i n g alpine marmots (Marmota mar mot a).. Anim. Behav. 24: 27-35. Barash, D. P. 1977. Sociobiology and Behavior. Else v i e r North-Holland Inc., New York, N.Y. 378 p. Beebe, F. L. 1974. F i e l d studies of the Falconiformes of B r i t i s h Columbia. Occas. Pap. B. C. Prov. Mus. 17: 1-163. Bent, A. C. 1937. L i f e H i s t o r i e s of North American Birds of Prey (Order Falconiformes - Part 1). D. S. Natl. Mus. Bu l l . 167, Washington, D. C. 409 p. Bibikov, D. I. 1967. Gornve-surki Srednei A z i i i Kazakhstana. Nauka, Moscow. Bopp, P. 1955. K o l o n i a l t e r r i t o r i e n bei Murmeltieren. Rev. Suisse Zool. 62: 295-299. 117 Bronson, F. H. 1964. Agonistic behaviour i n woodchucks. Anim. Behav. 12: 470-478. Brooke, R. C., E. B. Peterson, and V. J . Krajina. 1970. The SUbalpine Mountain Hemlock Zone. Subalpine vegetation i n southwestern B r i t i s h Columbia, i t s c l i m a t i c c h a r a c t e r i s t i c s , s o i l s , ecosystems and environmental relationships. Ecol. of Western No. Am. 2: 147-349. Brooks, S. J., and E.M. Banks. 1973. Behavioural biology of the coll a r e d lemming rDicrostpnyx groenlandicus ( T r a i l l ) ] : An ' a n a l y s i s of acoustical communication. Anim. Behav. Monogr. ;v 6: 1-83. , Brown, J. L. 1975. The Evolution of Behavior. W. W. Norton, New York, N.Y. 761 p. Brown, J . L., and G. Orians. 1970. Spacing patterns i n mobile animals. Annu. Rev. Ecol. Syst. 1: 239-262. Ca r l , G. C. 1944. The Vancouver Island marmot. V i c t o r i a Naturalist 1: 77-78. Charnov, E. L., and J. R. Krebs. The evolution of alarm c a l l s : Altruism or manipulation? Am. Nat. 109: 107-112. Cooper, W. S. 1942. An is o l a t e d colony of plants on a gl a c i e r clad mountain. B u l l . Torrey Bot. Club 69: 429-433. Cowan, I. McT., and C. J. Guiguet. 1965. The Mammals of B r i t i s h Columbia. B. C. Prov. Mus. Handbook No 11. 414 p. Crook, J. H. 1970. So c i a l organization and the environment: aspects of contemporary s o c i a l ethology. Anim. Behav. 18: ' 197-209. Davis, D. E. 1966. The molt of woodchucks (Marmota mpnax) . Mammalia 30: 640-644. Eisenberg, J . F. 1974. The functional and motivational basis of histricomorph vocalizations. Symp. Zool. Soc. Lond. 34: 211-247. 118 Emlen, J. M. 1973. Ecology: An Evolutionary Approach. Addison-welsey Publishing House, Reading, Mass. 492 p., Ewer, R. F. 1968. Ethology of Mammals. Paul Elek Ltd., London. 417 p. F a r r e l l , R. K., L. Koger, and B. Winward. 1966. Freeze branding of c a t t l e , dogs, and cats for i d e n t i f i c a t i o n . J. Am. Vet. Med. Assoc. 149: 745-752. F i t c h , H. S. 1948. Ecology of the C a l i f o r n i a ground s g u i r r e l on grazing lands. Am. Midi. Nat. 39: 513-596. F l i n t , R.F. 1971. G l a c i a l and Quaternary Geology. John Wiley and Sons Inc., New York, N.Y. 892 p. Foster, J. B., 1965. The evolution of the mammals of the Queen Charlotte Islands, B r i t i s h Columbia. Occas. Pap. B. C. •prov. Mus. 14: 1-130. Geist, V. 1971. Mountain Sheep: A Study in Behavior and Evolution. The University of Chicago Press, Chicago, 111. 383 p. George, W. 1974. Notes on the gundis (F. Ctenodactylidae). Symp. Zool. Soc. Lond. 34: 143-160. Gray, D. R. 1967. Behaviour and a c t i v i t y i n a colony of Hoary marmots (Marmota caligata) i n Manning Park, B. C. and a comparison of behaviour with other marmot species. B. Sc. Thesis, University of V i c t o r i a , V i c t o r i a , B.C. 91 p. Gray, D. R. 1975. The marmots of Spotted N e l l i e Ridge. Nat, Can. (Ottawa) 4(1): 3-8. Gregson, J. D. 1956. The Ixodoidea of Canada. (Ottawa) Science Service, Entomolgy Divis i o n , Canadian Dept. of Agriculture, Publ. #930. 92 p. Hadow, H. H. 197 2. Freeze-br anding: A permanent marking technigue for pigmented mammals. J . Wildl. Manage. 36: 645-649. 119 H a l p i n , Z . T . 1974 . I n d i v i d u a l d i f f e r e n c e s i n t h e b i o l o g i c a l o d o r s o f t h e M o n g o l i a n g e r b i l f M e r i o n e s u n q u i c u l a t u s ) . B e h a v . B i o l . 11: 2 5 3 - 2 5 9 . H a m i l t o n , W. D . 1964 . T h e g e n e t i c a l e v o l u t i o n o f s o c i a l b e h a v i o r . I a n d I I . J . T h e o r . B i o l . 7 : 1 - 5 2 . H a m i l t o n , W. H . , J r . 1934. T h e l i f e h i s t o r y o f t h e r u f e s c e n t w o o d c h u c k M a r m o t a monax r u f e s c e n s H o w e l l . A n n . C a r n e g i e M u s . 2 3 : 8 5 - 1 7 8 . H a r d y , G . A . 1 9 5 5 . T h e n a t u r a l h i s t o r y o f t h e F o r b i d d e n P l a t e a u a r e a on V a n c o u v e r I s l a n d , B r i t i s h C o l u m b i a . B. C . P r o v . M u s . N a t . H i s t . A n t h r o p o l . R e p o r t 1954: B 2 4 - B 6 3 . H a r i n g t o n , C . R . 1 9 7 5 . , P l e i s t o c e n e m u s k o x e n (Symbos) f r o m A l b e r t a a n d B r i t i s h C o l u m b i a . C a n . J . E a r t h S c i . 12 : 9 0 3 -9 1 9 . H a y e s , S . R . 1976. D a i l y a c t i v i t y a n d body t e m p e r a t u r e o f t h e s o u t h e r n w o o d c h u c k , M a r m o t a monax, i n n o r t h w e s t e r n A r k a n s a s . J . Mammal. 5 7 : 2 9 1 - 2 9 9 . H e u s s e r , C . J . 1954. N u n a t a k f l o r a o f t h e J u n e a u i c e f i e l d , A l a s k a . B u l l . T o r r e y B o t . C l u b . 8 1 : 2 3 6 - 2 5 0 . H e u s s e r , C J . 1960 . L a t e - P l e i s t o c e n e E n v i r o n m e n t s o f N o r t h P a c i f i c N o r t h A m e r i c a . A m e r i c a n G e o g r a p h i c a l S o c , New Y o r k , N . Y . 308 p . H o f f m a n n , R. S . 1974 . T e r r e s t r i a l v e r t e b r a t e s , p p . 4 7 5 - 5 6 8 . I n J . D . I v e s a n d R . G . B a r r y , e d . A r c t i c a n d a l p i n e e n v i r o n m e n t s . W i l l i a m C l o w e s and S o n s L t d . , L o n d o n . H o w e l l , A . H . 1 9 1 5 . R e v i s i o n o f t h e N o r t h A m e r i c a n m a r m o t s . N . ; Am. F a u n a 3 7 : 1 - 8 0 . I y e r s o n , S , L , a n d T u r n e r , B . N . 1 9 7 2 . N a t u r a l h i s t o r y o f a M a n i t o b a p o p u l a t i o n o f F r a n k l i n ' s g r o u n d s g u i r r e l s . C a n . F i e l d - N a t . 8 6 : 1 4 5 - 1 4 9 . 120 Ives, J. D. 1974. B i o l o g i c a l refugia and the nunatak hypothesis, :,; pj?.: . 605-636. In J. D. Ives and R. G. Barry, ed. A r c t i c and alpine environments. William Clowes and Sons Ltd., London. King, J. A. 1955. So c i a l behavior, s o c i a l organization, and population dynamics i n a biack-tailed p r a i r i e dog town i n the Black H i l l s of South Dakota. Contrib. Lab. Vertebr. B i o l . Univ. Mich. 67: 1-123. Koeing, V. L. 1957. Beobachtungen uber Reviermarkierung sowie, Droh-, Kampt-, und Abwehrverhalten des Murmeltiers (Marmota mar mot aL.). Z. Tierpsychol. 14: 510-521., Konishi, M. 1973. Locatable and nonlocatable acoustical signals . f o r barn owls. Am. Nat. 107: 775-7 85. Lazarus, A. B. , and F. P. Rowe. 1975. Freeze-marking rodents with a pressurized r e f r i g e r a n t . Mammal Rev. 5: 31-34. Lewontin, R. C. 1970. The units of selection. Annu. Rev. Ecol. Syst. 1: 1-18. Lloyd, J . E. . 1972. /Vocalizations i n Marmota monax. J. Mammal. 53: 214-216. Marler, P. 1955. Charact e r i s t i c s of some animal c a l l s . Nature 176: 6-8. Marler, P. 1957. S p e c i f i c d i s t i n c t i v e n e s s i n the communication signals of birds. Behaviour 11: 13-39. Mathews, L. H. 1971. The L i f e of Mammals. weidenfield and Nicolson, London. Vol 2. 440 p. , Mathews, W. , H., J . G. Fyles, and H. W. Nasmith. 1970. Postglacial c r u s t a l movements i n southwestern B r i t i s h Columbia and adjacent Washington state. Can. J. Earth S c i . 7: 690-702. Maynard Smith, J. 1965. The evolution of alarm c a l l s . Am. Nat. 99: 59-63. 121 Mayr, E. 1963. Animal Species and Evolution. Harvard University Press., Cambridge, Mass. 797 p. McCabe, T. T., and I.,McT. Cowan. 1945. . Peromy.scus maniculatus macrorhinus and the problem of i n s u l a r i t y . Trans. Royal Can. Inst. 25: 117-216. Melchior, H. R. 1971. Ch a r a c t e r i s t i c s of a r c t i c ground s q u i r r e l ! alarm c a l l s . Oecologia 7: 184-190. Murray, B. M., and D. F. Murray. 1969. Notes on mammals i n alpine areas on the northern St. E l i a s mountains, Yukon Ter r i t o r y and Alaska. Can. Field-Nat. 83: 331-338. Olendorff, R. R. 1976. Food habits of North American golden eagles. Am. Midi. Nat. 95: 230-236. Owings, D. H., M. Borchert, and R. V i r g i n i a . 1977. The behaviour of C a l i f o r n i a ground s q u i r r e l s . Anim. Behav. 25: 221-230. Pattie, D. L. 1967. Observations on an alpine population of yellow-bellied marmots f Marmota f l a v i v e n t r i s ) . Northwest Sci.,41: 96-102. Pearson, 0. P. 1948. L i f e history of mountain viscachas i n Peru. J . Mammal. 29: 345-374. Pianka, E. R. 1974. Evolutionary Ecology. Harper and Row, New ; York, N.Y. 356 p. Rausch, R. L., and V. R. Rausch. 1965. Cytogenetic evidence for the s p e c i f i c d i s t i n c t i o n of an Alaskan marmot, Marmota broweri H a l l and Gilmore {Mammalia: Sciuridae). Chromosoma 16? 618-623. Rausch, R. L., and V. R. Rausch. 1971. The somatic chromosomes of some North American marmots (Sciuridae), with remarks on the relationships of Marmota broweri Hall and Gilmore. Mammalia 35: 85-101. lopd, J . P. 1972. Ecological and behavioural comparisons of three species of Argentine cavies. Anim. Behav. Monogr. 5: 1-83. 122 Rowlands, I. ,W. 1974. Mountain viscacha. ,Symp. Zool. Soc. Lond. 34: 131-141., Shorey, H. H.,1976. Animal Communication by Pheromones. Academic Press, New York, N.Y. 167 p. Smith, H. J . , S. L. Smith, E. C. Oppenhiemer, and J . G. D e v i l l a . 1977. Vocalizations of the b l a c k - t a i l e d p r a i r i e dog, Cynomys ludovicianus. Anim. Behav. 25: 152-164. Snyder, B. L., D. E. Davis, and J . J . C h r i s t i a n . 1967.. Seasonal changes in the weights of woodchucks. J. Mammal. 42: 297-312. Stark, H. E. 1970. A r e v i s i o n of the f l e a genus Thrassis- Jordan 1933 (Siphonaptera:Ceratophyllidae). Univ. C a l i f . Publ. Ent. 53: 1-184. Steiner, A. L. 1975. "Greeting" behavior in some sciuridae, from ah : ontogenetic, evolutionary and socio-behavioral perspective. Nat. Can.(Que) 102: 737-751. Sterns, S. C. 1976. L i f e history t a c t i c s : A review of the ideas. Q. Rev. B i o l . 51: 3-47. Svoboda, J. 1972. Vascular plants productivity s t i d i e s of raised beach ridges (semi-polar desert) i n the Truelove Lowland, pp. 146-184. In L. C. B l i s s , ed. Devon Island I. B. P. Project, high A r c t i c ecosystem: project report 1970 and 1971. Dept. of Botany, University of Alberta. 413 p. Swarth, H. S. 1911. Two new species of marmots from northwestern America. Univ. C a l i f . Publ. Zool. 7: 201-204. Swarth, H. S. 1912. Report on a c o l l e c t i o n of birds and mammals from Vancouver Island. Univ. C a l i f . Publ. Zool, 10: 1-124. Taber, R. D., and I. McT. Cowan. 1971. Capturing and marking wild animals, pp. 277-317. In R. fl. G i l e s , ed. W i l d l i f e Investigation Technigues. 3rd ed. The W i l d l i f e Society, Washington, D. C. 123 Taulman, J. F. 1975. An ethological study of the hoary marmot, Marmota caligatacascadensis. M. Sc. Thesis, Central Washington State College, Ellensberg, Wash. 128p. Taylor, K. D. 1969. An anomalous freeze-branding re s u l t i n a rat. J. Zool. Lond. 158: 214-415. Tr i v e r s , R. L. 1971. The evolution of r e c i p r o c a l altruism. tQ. ;Rev. B i o l . 46: 35-57. Turner, L.' ,W.1973. Vocal and escape responses of Speraophilus ! beldingi to predators. J. Mammal. 54: 990-993. Vaughan, T. A., 1972. Mammalogy., W. B. Saunders Co., Philadelphia, Pa. 463 p. Vos, A. de, and I. G i l l e s p i e . 1960. A study of woodchucks on an Ontario farm. Can. Field-Nat. 74: 130-145. Walker, E. P., 3rd ed. 1975. Mammals of the World. The Johns Hopkins Universiyt Press, Baltimore, Md. 1500 p. Waring, G. H. 1966. Sound communication of the yellow-bellied marmot (Marmota f l a v i v e n t r i s ) . Anim. Behav. 14: 177-183. Waring, G. H. 1970. Sound communication of b l a c k - t a i l e d , white-t a i l e d and Gunnison«s p r a i r i e dogs. Am. Midi. Nat. 83; 167-185. Williams, G. C. 1966. Adaptation and Natural Selection. Princeton University Press, Princeton, N.J. 307 p. Wilson, E. 0. 1975. Sociobiology: The New Synthesis. Harvard University Press, Cambridge, Mass. 697 p. Wright, H. E., and D. G. Frey, ed. 1965., The Quaternary of United States. Princeton University Press, Princeton, N.J. 922 p. Wynne-Edwards, V. C. 1962. Animal Dispersion i n Relation to Social Behaviour. Oliver and Boyd, London. 653 p. 124 Yeaton, R. I. 1972. Social behaviour and s o c i a l organization i n Richardson* s ground s q u i r r e l Spermophilus r i c h a r d s o n i i i n Saskatchewan. J. Mammal. 53: 139-148. 125 Appendix I. A L i s t of Plant Species Found on the Haley Lake Study Area Indicating Those Species Known to be Eaten by Vancouver Island Marmots Family and S c i e n t i f i c Same Common Name Plants Eaten Selaginellaceae Selaginella wallacei Polypodiaceae Adiantutn pedatum Cryptogramma crispa Pply_stichum muni turn Pteridium aquilinum Cupressaceae Chamaecyparis nootkatensis Juniperis communis Pinaceae Abies amabilis Abies lasiocarpa Pinus conto rta Pinus monticpla Tsuga heterophylla Tsuga. mertensiana Pseudotsuga menziesii Betulaceae Alnus sinuata Aristolochiaceae *sarum caudatum Portulacaceae Clay_tonia lanceolata Caryophylaceae Arenaria macrop_hylla Silene menziesii s e l a g i n e l l a maidenhair fern parsley fern sword fern bracken fern yellow cedar dwarf juniper amabilis f i r alpine f i r lodgepole pine western white pine western hemlock mountain hemlock Douglas f i r Sitka alder wild ginger western springbeauty bigleaf sandwort ca t c h f l y fronds 0 bark° Ranunculaceae Actaea rubra M M i i S a i S . formosa Delphinium sp. . Thalictrum Occidentale banenerry columbine larkspur meadow rue lvs», f l w s 2 l v s , flws, fr' 3 126 Berberidaceae ASfeilus t r i g h y l l a Crassulaceae Sedum diyerqens Saxifraqaceae .;Saxif rag a bronchial i s ferruginea . .Saxifraga occidentalis ' Tellima grandiflorum Grossulariaceae Ribes lacustre v a n i l l a l e a f stonecrop spotted saxifraqe rusty saxifraqe western saxifrage fringecup swamp gooseberry Rosaceae f£a<jaria yirgin i a n a Luetkea pectinata P o t e n t i l i a d i y e r s i f o l i a Sorbus sitchensis blueleaf strawberry partridgefoot c i n q u e f o i l Sitka mountain ash Leguminosae Lathyrus neyadensis Lnjjj.nus l a t i f o l i u s Celastraceae Pachistima myrsinites Violaceae Viola g l a b e l l a Onagraceae Epilobium alpinum Ombelliferae Heracleum lanatum Lqmatium sp. Ericaceae Arctost aph y1gs uya-ursi Cassiope mertensiana Phyllodoce empetriform i s Ifeo^odendron albiflorum Vat eg in i u m c a e s gi t o s a Vaccinium deliciosum Vaccinium membraniceum laccinium ovalifolium l a S S i S i S S spp. Polemoniaceae Phlox d i f f u s a Labiatae ; liSSS111 vulgaris sweat-pea broadleaf lupine f a l s e box yellow v i o l e t alpine fireweed cow-parsnip b i s c u i t - r o o t kinnikinnick white moss-heather red heather white rhododendron dwarf huckelberry blue-leaf huckelberry t h i n - l e a f huckelberry oval-leaf huckelberry huckelberry spreading phlox s e l f - h e a l lvs l v s * , flws* l v s , f r * lvs° l v s , f r * l v s , flws* 127 Scrophulariaceae £astilleja miniata C a s t i l l e j a p a r y i f l o r a UiJBiiiiJS guttatus ge'dlcularis bracteosa Penstemon dayidsonii ' 2§I2SS£a wormsk-ioldii Rubiaceae Galium boreale Indian paintbrush Indian paintbrush yellow monkey flower lousewort penstemon speedwell northern bedstraw ns*, flws* ns, flws Caprifoliaceae Sambucus racemosa red elderberry bark Valerianaceae Valeriana s i t c h e n s i s lountain valerian lvs Campanulaceae Campanula r o t u n d i f o l i a Compositae Achillea millefolium Agosetis aurantiaca , Anajghalis margarjtacea Arnica l a t i f o l i a -Cirsium edule ' Erigjeron £eregrinus Efpphyllum lan a turn • Senicio t r i a n g u l a r i s Tar.axkcum o f f i c i n a l e b l u e b e l l flws yarrow agoseris pearly everlasting broad-leaf arnica Indian t h i s t l e lvs° mountain daisy woolly sunflower lvs° giant ragwort common dandelion Juncaceae Juncus drummondii rush Cyperaceae Carex mertensii Carex nigricans Carex s p e c t a b i l i s Carex spp.. Graminae Agrostis diegoensis Bromus sitc h e n s i s Elymus glaucus Mellcia sub a l a t a Phleum alpinum Trisetum spicatum Lilaceae Alium crenulatum E'rit'hrpnium grandiflorum Lillum eg1umbi anum Smilacina racemosa Stenarithium Occidentale T r i l l i u m ovaturn Veratruffl y i r i d e sedge sedge sedge sedge bentgrass brome grass blue wild-rye Alaska oniongrass mountain timothy spike trisetum wild onion avalanche l i l y t i g e r l i l y f a l s e Solomon*s seal western stenanthium western t r i I l i u m f a l s e hellebore l e a f t i p s , ns l v s , ns. l v s , ns, flws l v s , flws* lvs 128 Orchidaceae Habenaria sp. bog orchid Lichens Cladonia s p . Pe l t i g e r a a p tho s a So l o r i n a c r o c e a V s t e r e o c a n l o n s p . Thanolia • s p . ., Umbilicaria s p . , r o c k t r i p e t h a l l i * leaves z flowers 3 f r u i t * new shoots * preferred food ° rarely eaten CM Appendix I I . A L i s t o f a l l Known C o l l e c t e d , T o t a l = 30 V a n c o u v e r I s l a n d Marmot Specimens t h a t have been Year C o l l e c t o r C o l l e c t e d C o l l e c t i o n Localities*»> Number C o l l e c t e d P r e s e n t L o c a t i o n of Specimens B. S. Swarth 1910 A. Peak ' 1929 D. I . Walker 1930 ft. Pacey and . 1931 I. tel. Cowan P. S. M a r t i n 1938 1. E. G a t e n l y 19U0 P. L. Deebe D. G. King 19U3 1965 P. L. B r i g h t 1968 J . Csman D. C. Heard 1968 1968 ? 1971 Mt. D o u q l a s K i n g Solomon E a s i n G o l d e n E a g l e E a s i n E a t t l e d t . ( l o c a t i o n unknown) J o r d a n Meadows Oruyn Mountain colony one Ht. A r r o w s m i t h D r i n k Water C r e e k Mt. Washington Mt. Washington Rt. Washington Comox B e a u f o r t Range Green M o u n t a i n c o l o n y one 7 ca) 3 1 1 1 ( 3 > 1 CJ> 1 1<*> 1 1 1 < 3 > 1 < 3 > 1 ( 3 ) U n i v e r s i t y o f C a l i f o r n i a , B e r k l e y , n u t t e r s 12090-12100. N a t i o n a l Museum o f Canada, Ottawa, number 10333. B r i t i s h Columbia P r o v i n c i a l Museum, r II IT h o r 12 f> 0. rin Ivor::! t y o f BrU. l;ih C o l u m b i a , n u n t o r s 1-S8f»fi and »92B (n = 7) , 6 N a t i o n a l Museum o f Canada, Cttawa, numbers 1«088-m089 (n = 2) . B r i t i s h Columbia P r o v i n c i a l Museum, nu n t e r 2898. B r i t i s h Columbia P r o v i n c i a l Museum, nurcter U5U0. B r i t i s h Columbia P r o v i n c i a l Museum, nu n t e r 5021. O n i v e r s i t y o f A l a s k a , n u n t e r 28751. U n i v e r s i t y o f Montana, n u n t e r UMZM 13521. E r i t i s h Columbia P r o v i n c i a l Museum, un <ataloqued. E r i t i s h Columbia F i s h and W i l d l i f e B r a n c h , P o r t A l b e r n i O f f i c e O 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 , unca t a l o q u e d . 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 , t e a c h i n g c o l l e c t i o n . <o mapped cn F i g u r e 1; c o o r d i n a t e s i n T a b l e I <«> i n c l u d e s tie t y p e s p e c i m e n w i t h the c c l m c l e t e s k e l e t o n < 3 > s k u l l c r l y t . ,, . . t o s k i n o n l y , may go w i t h t h e s k u l l i n the DEC t e a c h i n g c o l l e c t i o n ^ 

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