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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  MARMOTA  HABBOTS,  VANCOUVBRENSIS  by DOUGLAS C . E.Sc., A THESIS THE  University  SUBMITTED  HEARD  of  Waterloo,  1973  IN  PARTIAL  PULPILLMENT  REQUIREMENTS  FOR THE  DEGREE OF  MASTER OF  OF  SCIENCE  in THE F A C U L T Y (The  OF GRADUATE  Department  We a c c e p t t h i s  thesis  reguired  THE  UNIVERSITY  Zoology)  as conforming t o standard  OF B R I T I S H  November,  Douglas  of  C.  STUDIES  COLUMBIA  1977  Heard,  1977  the  In  presenting  an  advanced degree  the I  Library  further  for  shall  agree  scholarly  by  his  of  this  thesis at make  that  thesis  partial  it  freely  permission  for  for  financial  is  University  of  British  ?  ^2^07^  of  Columbia,  British  by  gain  Columbia  for  the  understood  of  2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5  of  extensive  p u r p o s e s may be g r a n t e d It  fulfilment  available  permission.  Department  Date  in  the U n i v e r s i t y  representatives.  written  The  this  shall  requirements  reference copying  Head o f  that  not  the  of  I  agree  and this  be a l l o w e d  or  that  study. thesis  my D e p a r t m e n t  copying  for  or  publication  w i t h o u t my  ABSTRACT  I studied the s o c i a l marmot, 197*».  Marraota  at  of  the  Vancouver  Island  V a n c o u v e r e n s i s , d u r i n g the summers of 1973 and  V i r t u a l l y nothing  species  behaviour  the  was known about t h e behaviour  outset  of  t h i s study,  Barash  suggested t h a t t h e s o c i a l behaviour and s o c i a l  of  this  {1973b, 1974a)  organization  of  marmot s p e c i e s was determined by the s e v e r i t y o f the environment (the  vegetative  r a t e of marmots. short  growing  growing  The  hypothesis  and i t s e f f e c t  He p r e d i c t e d t h a t season  marmot  environments  that s o c i a l t o l e r a n c e increased.  season)  would  by observing  of  species  living  in  would be h i g h l y s o c i a l but  decrease  objective  on the growth  this  as  the  study  growing  was  to  season  test  this  t h e s o c i a l behaviour of Vancouver I s l a n d  marmots and comparing t h i s  to  the  length  of  the  vegetative  growing season.  J3« vancouyerensis Columbia. crossed  is  The o r i g i n a l to  Vancouver  endemic t o Vancouver I s l a n d , B r i t i s h  colonizers  of  this  species  I s l a n d v i a land connections  probably  that existed  during the I l l i n o i a n g l a c i a l p e r i o d , approximately 100,000 ago,  and s u r v i v e d subseguent refugia  isolated  from mainland forms f o r a length years)  adaptations  both.  maxima  coastal  100,000  or  glacial  sufficient  to t h e i r  on  years  nunataks  and  Vancouver I s l a n d marmots have been  to  show  of  time  specific  (10,000  to  evolutionary  Vancouver I s l a n d environment.  Vancouver I s l a n d marmots l i v e  in  small  colonies  i n the  iii  subalpine one  parkland.  adult  S o c i a l groups c o n s i s t e d o f one  female,  yearlings,  and  and  infants.  with a l a r g e amount members.  Alarm  predators  and  whistles  could be  locate. colony  long  predators  numbers  of  two-year-olds,  S o c i a l groups were h i g h l y communication were  given  heard  given  (e.g. eagles) and  bandwidth  of  calls  were  terrestrial  variable  a d u l t male,  in  over  in  occurring  whole  response  whistles  to  were  (e.g. black  among  response  the  to potential  aerial in  predators  response  was  a nose t o u c h i n g  occurred  behaviour termed  of  greetings  as other s o c i a l behaviour p a t t e r n s  as  well  Vancouver  that  A l l age-sex c l a s s e s  Island  vegetative marmots  growing was  season  marmots  most  closely  Barash*s  experienced  approximately  much  hypothesis  the  behaviour  resembled  where the growing season i s rejected  Island  same  of  M. oly_i£us, shorter.  engaged  growing season not taken  by  in  i n about  On  this  l e n g t h of time r e g u i r e d  to r e a c h  living  basis  maturity.  correlated  I  vegetative that  vegetative but  measured d i r e c t l y .  positively  of  Island  variability  be used as an index of growth r a t e  degree of s o c i a l t o l e r a n c e i s  that  a species  I suggest t h a t  to reach a d u l t s i z e be  Vancouver  as  t h a t the l e n g t h of the  Barash observed among marmot s p e c i e s .  time  among  greeting.  Vancouver  growing season i s s u f f i c i e n t t o account f o r the  the  to  proportions.  M . , f l a v i v e n t r i s but the s o c i a l marmots  to  Both c a l l s are narrow  most common s o c i a l behaviour  members  The  Short  sounds, a c h a r a c t e r i s t i c t h a t makes them d i f f i c u l t The  the same  colony  colony.  given  bears).  integrated  with  that The the  iv  TABLE Abstract  OF  CONTENTS  ................. .................................  List  of  Figures  List  of Tables  vi viii  Acknowledgements  ..........................................  Introduction  ..  Sociobiology Vancouver  and  and  ..  Barash's  Island  Taxonomy  Hypothesis  ...................  Marmots Evolutionary  History  ...................  5  ...  10 10  Observation  Analysis  of  Trapping  and  Measurement  Social  10  Behaviour  Data  ......................  Marking of  Vocalizations Results  Microclimate  ............................  21  ..........................................  21  Colony  22  Characteristics  Physical  22  Characteristics  26  Composition  Activity  Patterns  Vocalizations  14 19  ........................ . .  Habitat  1 4  Areas  Methods o f  x 1  Methods Study  i i  33 ......................................  .... ...  ......  34 44  Whistles............................  44  Keeaws  54  Rapid  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v .,... . . , . Chirps  Hisses Tooth  ........................................  57  ..............................................  57  Chatters  Screams  and  Growls  57 ..................................  59  S o c i a l Behaviour  59  S o c i a l Behaviour P a t t e r n s  59  Dominance R e l a t i o n s h i p s  63  The Frequency o f S o c i a l Behaviour P a t t e r n s  ..........  66  I n t e r a c t i o n Sequences ...............................  72  Rates of S o c i a l Behaviour ...........................  74  Dispersion, T e r r i t o r i a l i t y ,  and Scent Harking .......  82  ................ *...............................  91  v o c a l i z a t i o n s ..........................................  91  Discussion  interspecific  Comparisons  o f Marmot V o c a l i z a t i o n s ..  A l t r u i s m and Marmot Alarm C a l l i n g  91  ...................  94  The E v o l u t i o n o f H h i s t l e S t r u c t u r e ..................  97  The E v o l u t i o n o f W h i s t l e Function  99  I n t e r s p e c i f i c Comparisons  ...................  o f Marmot S o c i a l Behaviour . . 1 0 2  A T e s t o f Barash's Hypothesis ....................... 102 L i t e r a t u r e C i t e d .............. ................ ....,....... 115 Appendix Lake  I . A L i s t of Plant Species Found Study  Area  Indicating  on  the  Haley  Those Species Known t o be  Eaten by Vancouver I s l a n d Marmots ....................... 125 Appendix I I . A L i s t Specimens  that  of have  a l l Known Vancouver I s l a n d  Marmot  been C o l l e c t e d , T o t a l = 30 ...... 129  vi  LIST OF FIGURES Figure  1. L o c a t i o n s o f  Known  Colonies  Vancouver  Island  Marmot  .  11  Figure  2. Photographs of t h e Haley Lake Study Area ....... 12  Figure  3. Photographs o f Vancouver I s l a n d Marmot H a b i t a t  Figure  4. Seasonal Changes  in  the Weights  of  .23  Vancouver  I s l a n d Marmots . . . . . , Figure  ;  27  5. Seasonal Changes i n t h e Weights of Adult Marmots  ..... . .. .  . . ..... .. . ... . . .......  .  . .....  .... 29  Figure  6. A c t i v i t y  Budgets f o r May, a l l Animals Combined . 3 9  Figure  7. A c t i v i t y  Budgets f o r June, a l l Animals Combined  40  Figure  8. A c t i v i t y Budgets f o r J u l y , a l l Animals Combined  41  Figure  9.  Activity  Budgets f o r  August, a l l  Animals  Combined ........................... .................... 42 F i g u r e 10.  Activity  Budgets f o r September,  all  Animals  Combined  43  F i g u r e 11. R e p r e s e n t a t i v e Sonogram of a Short Whistle ..... 45 F i g u r e 12. R e p r e s e n t a t i v e Sonogram of a Long W h i s t l e ...... 46 F i g u r e 13.  A  Comparison of the  Length o f  Long and Short  W h i s t l i n g Sequences ...... .......... .................... 48 F i g u r e 14. R e p r e s e n t a t i v e Sonograms o f Keeaws F i g u r e 15. Occurred  Freguency with which Together and  55  Whistles and  Separately  With and  Keeaws  Without  a  Known Disturbance as a Stimulus ........................ 56 F i g u r e 16. R e p r e s e n t a t i v e Sonogram o f a H i s s .............. 58 F i g u r e 17. Temporal Context With  Which  Two-act  Sequences  Occurred ......... ... . .............. . . . . . . . . . . . . . . . . i . . . F i g u r e 18.  A  Comparison  Of  Interaction  Rates  Between  73  C o l o n i e s One and Two ................................... 75 F i g u r e 19. G r e e t i n g and  agonistic  Interaction  Rates per  Dyad . ..... ...... ..... . .. . ..... ........ ................. 79 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 Rates Among Months .... 80 F i g u r e 21.  V a r i a t i o n i n the G r e e t i n g  Rate Throughout the  Day . ,  81  F i g u r e 22.  Home  Ranges o f the  Females (#12 and #15) i n May Males (#13 and #17) i n June  1974, 1974, on  Two  Adult  and the Two Adult The  Haley  Lake  Study Area ..... .................... .... ................ 83 F i g u r e 23. Home Ranges o f Four Female Marmots on t h e Haley Lake Study Area i n J u l y 1974  84  F i g u r e 24. Home Ranges of the Two Adult Males on the Haley Lake Study Area i n August 1974 ......................... 85 F i g u r e 25.  Rates o f  Scent  Marking by  Adult  Sales  #4,  #13, and #17 ........................................... 88 F i g u r e 26. A  Comparison o f  Greeting  Species F i g u r e 27. Species  Rates  Among  Marmot  ............ ... .....105 A  Comparison o f  chasing  Rates  Among  Marmot  .................................... ............107  F i g u r e 28. A Comparison o f t h e R a t i o of G r e e t i n g s Among, Marmot Species  t o Chases  ...................................108  LIST OF TABLES Table  I . Habitat  Characteristics  of  Vancouver  Island  Marmot C o l o n i e s ........................................ Table  I I . Climatic  Study (  Area  Characteristics and  L o c a t i o n i n the  a  Coast  of  Typical  the  Haley Lake  Parkland  Mountains,  after  Subzone  Brooke e t a l •  1970 ........... Table I I I .  Age and  One and Two Table Table  13  25 sex  Composition o f  Marmot  Colonies  ............................................35  IV. Causes of W h i s t l i n g and Keeaw Sequences ........ 50 V.  Dominance  Matrices of  Age and  Sex C l a s s e s of  Vancouver I s l a n d Marmots ...............................,64 Table  VI. The  Frequency  Of  Occurrence  Of  Each  Social  Behaviour P a t t e r n Between Age-sex C l a s s e s Table VII.  Percentages o f  Social  67  Behaviour  P a t t e r n s per  Age-sex C l a s s Table V I I I .  68 Comparisons of the  Frequency with which  Relative  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 ...................... Table  IX.  Interaction  Rate per  Age-sex  C l a s s of  B e h a v i o u r a l Acts per Thousand Hours .................... Table  X.  Interaction  Rate per  Dyad per  70  76  Behaviour per  Thousand Hours ................ .................. ....... 77 Table  XI. A Comparison of V o c a l i z a t i o n s w i t h i n the Marmota  Caligata. Group ..................... .................... Table XII. A L i s t Of Mammalian  Species  That  Have  92  Narrow  Bandwidth Alarm C a l l s ..................................101 Table X I I I .  &  Comparison o f t h e  R e l a t i v e Growth Rates Of  Yearling  Marmots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ........110  X  ACKNOWLEDGEMENTS  I  would  like  to  s u p e r v i s i o n , encouragement, of t h i s  I  thank  Dr. I . McT. Cowan  and p a t i e n c e throughout  for  his  the course  project.  would a l s o l i k e t o acknowledge  the help I r e c e i v e d  Ted Barsby, B r i t i s h Columbia F i s h and W i l d l i f e Branch  from:  (Nanaimo),  Dr. Fred B u n n e l l , J u d i t h Donaldson, Jack Evans, Dr. N. R. L i l e y , MacMillan B l o e d e l Floy  (Chemainus D i v i s i o n ) , Dr. J . Mary  Zitten.  During  this  study  I was supported by a N a t i o n a l Research  C o u n c i l of Canada S c h o l a r s h i p and Department  of Zoology Teaching  A s s i s t a n t s h i p s . My r e s e a r c h was f i n a n c e d by the Fund  T a y l o r , and  (Canada).  World  Wildlife  1  INTRODUCTION  SOCIOBIOLOGY AND  BARASH'S HYPOTHESIS  R e c e n t l y , t h e r e has been a marked i n c r e a s e i n the number o f investigators  who  natural selection Brown 1975, ecology  are  considering  (Crook 1970,  Wilson 1975).  (Crook 1970)  behaviour  Barash  as a product o f  1974a S 1977,  Alcock  S o c i o b i o l o g y (Wilson 1975)  or  1975, socio-  i s the s y s t e m a t i c study of the e v o l u t i o n o f  s o c i a l behaviour and s o c i a l o r g a n i z a t i o n i n r e l a t i o n t o ecology, demography,  and  population genetics.  T h i s new  approach t o the  study of animal behaviour has l e d t o the formation hypotheses concerning the a d a p t i v e s i g n i f i c a n c e fitness),  of  study was  social  behaviour  testable  ( c o n t r i b u t i o n to  and s o c i a l o r g a n i z a t i o n .  (1973b, 1974a) put f o r t h a hypothesis t o account f o r  the e v o l u t i o n of marmot s o c i e t i e s differences correlated of  in  social  behaviour  (Marjtota spp) . among  marmot  with v a r i a t i o n s i n the environment, the  He showed that  v e g e t a t i v e growing season.  species  snow  cover  (Barash  i n h a b i t environments growing animals burrows  seasons.  of They  (Bronson 1964) when  1973b). low  Barash*s measure o f  Woodchucks  elevation  are s o l i t a r y  were  s p e c i f i c a l l y , the  growing season was the number o f f r o s t - f r e e days i n the of  This  an attempt to t e s t one such h y p o t h e s i s .  Barash  length  of  with  absence  (Warmota long  monax)  (150  day)  and r e l a t i v e l y a g g r e s s i v e  and the young d i s p e r s e from  weaned (Vos and G i l l e s p i e 1960).  their  Olympic  (H. olyrnpus) i n h a b i t high e l e v a t i o n a l p i n e meadows that  natal marmots have  a  2  very  short  (40  to  70 day) growing season.  They l i v e i n w e l l  i n t e g r a t e d c o l o n i e s and are s o c i a l l y t o l e r a n t , social  behaviour  greetings until  i s characterized  (Barash 1973b, 1971a).  their  fourth  Barash's hypothesis  summer  by  a  that  high  i s , their  freguency  Olympic marmots do n o t  and  disperse  of  mature  during t h e i r  third.  i s t h a t s o c i a l t o l e r a n c e , as measured by the  r a t e with which g r e e t i n g s a r e performed, i s i n v e r s e l y r e l a t e d t o the length o f t h e v e g e t a t i v e growing season.  He  reasoned  that  1) s h o r t e r growing seasons r e s u l t i n lower growth r a t e s i n terms of  the  time  reguired  to  disadvantageous f o r subadult still  "undersize",  where  reach  adult  species;  3) i n  d i s p e r s e r s should  size  is  measured  be e i t h e r the same  inhabiting  4) a g g r e s s i o n the  they  are  as the weight o f an  adult  of the  proportion  of  the  adult  be r e l a t i v e l y more mature i n  increasingly  severe  environments;  from a d u l t s causes the young to d i s p e r s e ; thus  increase  progressively increasing  i t is  a l l marmot s p e c i e s , the minimum s i z e f o r  weight o r d i s p e r s i n g marmots should species  2)  marmots to d i s p e r s e when  d i s p e r s i n g marmots r e l a t i v e t o the weight of same  size;  in  social  shorter  necessity  tolerance  growing to  among marmots e x p e r i e n c i n g  seasons  inhibit  5)  the  may  be  due  to  the  d i s p e r s a l o f undersized  animals.  He  also  suggested  that  1)  i t i s important  population  s i z e more c l o s e l y r e g u l a t e d  increases  because  and  2)  have  as environmental s e v e r i t y  under severe c o n d i t i o n s the h a b i t a t would be  e a s i l y overgrazed, and t h e r e would food;  to  be  strong  competition  i f marmot s o c i a l behaviour r e g u l a t e s  for  population  3  s i z e i n a d e n s i t y dependent way; then 3) s o c i a l t o l e r a n c e should i n c r e a s e i n severe environments because the o p e r a t i o n of such system  a  of population r e g u l a t i o n would r e q u i r e a c l o s e r p h y s i c a l  proximity o f colony members.  Barash*s h y p o t h e s i s i s t e s t a b l e i n degree  of  social  tolerance  for  any  that  i t predicts  the  marmot s p e c i e s when the  l e n g t h of the v e g e t a t i v e growing season under which t h a t s p e c i e s evolved i s known. observing  the  social  (M. f / l a v i v e n t r i s , Barash  1974b),  behaviour  the  European  His observations  the y e l l o w - b e l l i e d  alpine  of  hoary  marmot  (B. c a l i q a t a ,  marmot and  by  (M. marmota, yellow-bellied  were c o n s i s t e n t with h i s hypothesis, but a l p i n e marmots  engaged i n s i g n i f i c a n t l y the  of  Barash 1973a) , the hoary marmot and  Barash 1976b) . marmots  Barash s e t out to t e s t h i s own h y p o t h e s i s  basis  of  the  fewer g r e e t i n g s than were  result.  ( s o c i a l t o l e r a n c e ) , may be i n a p p r o p r i a t e  obscures  the  1976b).  differences chasing,  based  Downhower,  and  on  between  which  Unfortunately,  hypothesis  of  this  that  it  d i s c r e t e parameters such as vary  independently  (Barash  he does not go on t o develop a r e v i s e d  these  Svendsen  may  in  discrete  (1976)  and  parameters. Anderson,  Armitage, Armitage, and  Hoffmann (1976) attempted to r e j e c t Barash*s hypothesis yellow-bellied  one  He suggested that a blanket term such as  sociality  g r e e t i n g and  on  s h o r t growing season o f i t s a l p i n e h a b i t a t .  Barash d i d not r e j e c t h i s h y p o t h e s i s on the b a s i s inconsistent  expected  marmots l i v i n g  because  at high e l e v a t i o n s grew f a s t e r i n  terms of grams per day than d i d y e l l o w - b e l l i e d marmots l i v i n g a t lower e l e v a t i o n s .  However, t h i s does not c o n s t i t u t e a  test  of  4  Barash's  h y p o t h e s i s s i n c e h i s hypothesis i s based on the growth  r a t e i n terms of the time taken t o reach a d u l t s i z e . taken  time  t o reach a d u l t s i z e i s not r e l a t e d s o l e l y t o the a b s o l u t e  weight gained per day, but i t i s also a f u n c t i o n of of  The  time  each  absolute  year  weight  of  during  which  adult  marmots  the  length  marmots gain weight and the of  the  species  being  considered.  At  the  two marmot studied,  outset  species  o f t h i s i n v e s t i g a t i o n there s t i l l in  North  M. vancouverensis  America  and  had  M. broweri.  t h i s study was t o document t h e l i f e  history  Vancouver I s l a n d marmots and t o provide hypothesis of marmot  which  remained  never  been  The o b j e c t i v e o f and  behaviour  of  another t e s t of Barash's  sociality.  VANCOUVER ISLAND MARMOTS  The 1911,  Vancouver I s l a n d marmot, Marmota yancouyerensis Swarth  i s endemic t o Vancouver I s l a n d , B r i t i s h Columbia.  in  small  colonies  on  steeply  sloping  Vancouver I s l a n d marmots are a c t i v e f o r only summer.  To  hibernate  f o r about e i g h t months o f t h e year.  be  avoid  subalpine a few  Living meadows,  months  the r i g o r s of the mountain winters,  each  marmots  Even then i t  may  necessary f o r them t o burrow out through many metres of snow  i n the s p r i n g .  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 I s l a n d marmots. was 1944,  based on three very  brief reports  This  information  (Swarth 1912: 89-90, C a r l  Hardy 1955: B61) which i n d i c a t e d that the n a t u r a l  history  5  of  Vancouver  Island  marmots  a l p i n e marmots (e.g.  was  s i m i l a r to other  species of  Barash 1973b) and amenable to study  using  s i m i l a r methods.  Taxonomy and  Evolutionary  Swarth  described  s p e c i e s i n 1911.  The  M. -Vancouverensis species  deeply  quite  Island  external  different  Howell  marmots M.  1915  and,  Howell on  2) the  and  PT. broweri  association  for  morphological fill  (Siphonaptera) were  obvious  dark  to  M. c a l i g a t a In  be  This  1965,  group  appears  Their ectoparasites within  collected  M.  identified  from by  Mr.  G.  in  the  reflect  this  the  natural  the  purely  M. c a l i g a t a  Holland  the  group.  vancouverensis P.  Rausch  within a  as  with  have s i m i l a r behaviour  also  relationships  be  well  Species  and  species to  as  included along  Rausch  separate  reasons  He  group  a  biological  is  Rausch 1971).  similarities.  s i m i l a r e c o l o g i c a l niches and  evolutionary  most  three groups of North American  ones used by Howell.  (see d i s c u s i o n ) .  study  the  M. olympus.  group.  group  designated  in  i!« c a l i g a t a  of  other marmot  pelage which i s uniformly  morphological  vancouverensis  considered  new  However, the karyotype of M. v a n c o u v e r e n s i s  based  M« . c a l i g a t a  any  The  very s i m i l a r to that of M. c a l i g a t a (Rausch and  In  as a  characteristics  from  1915).  marmot  1) the p o s t e r i o r border of the nasals which  V-shaped  brown to b l a c k .  Vancouver  c r a n i a l and  1911,  c h a r a c t e r i s t i c s are: is  the  are  (Swarth  History  during  close Fleas this  of A g r i c u l t u r e  6  Canada as T h r a s s i s ( T h r a s s i s ) s p e n c e r i subspecies  of  f l e a i s found  group (Stark 1970). from  Since  from any  ticks,  I  other  like  am  unaware  species  fleas,  Wagner.  (Acarina: of  Ixodidea)  t i c k s having  i n the M. c a l i g a t a  tend  to  may  represent  a  be host s p e c i f i c  new  Ixodes (P. Zuk, Canadian Department communication).  Since  the s p e c i e s probably  (Gregson  i n the genus  of Agriculture,  only one subadult  probably  M. c a l i g a t a  personal  specimen i s a v a i l a b l e  began  to  diverge  from  and becoming i s o l a t e d t h e r e .  fact  mammal  cavifrons, and  the f o l l o w i n g  species:  musk  ox  mastodon Mammut americanum, mammoths Mammut  Mammut cjolumbi, horse  Vancouver  Island  the  stock a f t e r c r o s s i n g to Vancouver I s l a n d  on temporary l a n d connections that  from  cannot be d e s c r i b e d .  J» i§2co«verensis ancestral  species  been  group.  1956), i t was not s u r p r i s i n g to d i s c o v e r t h a t t h e specimen M. vancouverensis  This  o n l y on marmots i n the M. c a l i j a t a  I c o l l e c t e d one t i c k  M.vancouverensis.  collected  spenceri  existed  Harington  (1975) b e l i e v e s  Symbos  imperator  Eguus sp., and Bison sp. once l i v e d on  suggests  mainland  The  that  land  connections  d u r i n g the l a t e P l e i s t o c e n e that  connections  with  (Harington between  the 1975).  Vancouver  I s l a n d and the mainland e x i s t e d on two occasions d u r i n g the l a s t (Fraser/Wisconsin) penultimate with  (Illinoian)  the mainland  maximum  about  depressed  and  glaciation.  probably  development  glaciation was  glaciation  of  at  least  once  during the  The most r e c e n t  connection  e x i s t e d j u s t p r i o r t o the time o f  continental  20,000 years ago.  i c e during  the  Fraser  At t h i s time the sea l e v e l  about 120 m below t h e present  sea l e v e l  and a  7  narrow c o r r i d o r would have j o i n e d Port Angeles and 1).  During  the  Peak  100,000 years  ago,  estimated  be  to  of  the  Illinoian  Wright  and  Frey  160  c o r r i d o r would have j o i n e d Vancouver Marmots  could  also  have  deposits that f i l l e d during  crossed  glaciation  196 5)  m below present  Victoria  sea  on  to  (roughly  levels  l e v e l s and  Island  were  an even wider the  mainland.  the massive f l o o d p l a i n  the whole S t r a i t o f Georgia r e g i o n  (Fig  the Olympia I n t e r g l a c i a t i o n about 30,000 years ago  from F l i n t  (Fig  1)  (date  1971) .  Although speciation,  little Mayr  is  (1963)  known has  about  the  estimated  rate  that  of  even  animal a rapidly  e v o l v i n g i s l a n d form would r e q u i r e a minimum of 100,000 years to achieve f u l l s p e c i f i c s t a t u s .  I f Mayr i s c o r r e c t ,  likely  Vancouver I s l a n d during  t h a t marmots c o l o n i z e d  the I l l i n o i a n  glaciation  rather  than  at  it  either  is or  most before  opportunity  during the F r a s e r g l a c i a t i o n .  The  Vashon  Vancouver I s l a n d if  Stade of the  (Mathews, F y l e s , and Nasmith 1970).  marmots c r o s s e d  to Vancouver I s l a n d before  must have been e i t h e r nunataks above  the  glaciers)  marmots could nunataks and  There  are  which are vegetated  t h i s period,  (high peaks and  ridges  there  emerging  G e o l o g i c a l evidence i n d i c a t e s t h a t  (Heusser 1960, (W.  Therefore,  or c o a s t a l r e f u g i a a v a i l a b l e on which the  survive.  coastal refugia  existed.  Fraser g l a c i a t i o n covered most of  Mathews et a l 1970, H.  Mathews  Muller  personal  e x i s t i n g nunataks i n Alaska  (Cooper  19 42,  Heusser  1954)  both  unpublished)  communication) and and  the Yukon support  8  arctic  ground  {Hurray and species  squirrel  (SoerraoDhilus  Hurray 1969) .  of  mammals  the Queen C h a r l o t t e  Foster  undulatus)  (1965)  populations  concluded  that  some  s u r v i v e d the l a s t g l a c i a t i o n on r e f u g i a on Islands.  Thus  it  is  not  difficult  to  imagine Vancouver I s l a n d marmots s u r v i v i n g the Vashon g l a c i a c i o n on s i m i l a r r e f u g i a . refugia  Other z o o l o g i c a l evidence a l s o supports the  concept.  No  species  of  mammal  vancouverensis p r e s e n t l y e x i s t s e x c l u s i v e l y subalpine  environment  s p e c i e s on the  island  recognized  a  ptarmigan  as  in  Vancouver I s l a n d and  and  against the  Ives  species.  In a d d i t i o n , w h i t e - t a i l e d  a  are  well  also  point  (HcCabe and  Cowan 1945).  nunatak s u r v i v a l  being  in  alpine  race  (L.  leucurus  is  confined  to  Other arguments f o r  hypothesis  are  discussed  by  (1974).  land connections p e r i o d and  that  existed  during  on  nunataks  or  coastal  g l a c i e r s r e t r e a t e d f r e e d i s p e r s a l was existence  of  the  Illinoian  glacial  would  have  peaks that they now colonization  of  refugia  or both.  Fraser As  the  probably made p o s s i b l e  by  a l p i n e h a b i t a t at the edge of the r e t r e a t i n g  As the f o r e s t s  marmots  to Vancouver I s l a n d v i a  the s p e c i e s s u r v i v e d the g l a c i a l maxima of the  glaciation  ice.  of  found  differentiated  ..'..•!!•'-K-2Sil£2S2§ESS§i§- probably crossed  the  no other mammal  distinct  and  Island  alpine-  to  s S ^ a t a l i s ) of t h i s r e l a t i v e l y i m p l a s t i c s p e c i e s Vancouver  the  diverged  leucurus)  the  than  has  {Lagop_us  environments  on  other  closed  in  gradually occupy.  mountain  below  them  Vancouver  become i s o l a t e d on the  A s i m i l a r example of tops  has  been  Island mountain  post  glacial  postulated  f o r the  mountain hare (Leo us t i m i d us) i n Europe  (Hoffmann  1974)  10  METHODS  STUDY AREAS  I made most of my  observations  at  two  about one k i l o m e t r e apart on the southern on  Vancouver I s l a n d , B r i t i s h Columbia  The  colony on the Haley Lake study area  and  2)  first  has  presumably  d i s c o v e r y i n 1932  characteristics  have  communication).  I  colonies  end of Green Mountain,  (numbers 1 and  by K. Racey and  also  changed made  2, F i g 1).  (colony number 1, F i g  been i n continuous  not  situated  existence since i t s  I . McT. ( I . McT.  Cowan, s i n c e i t s Cowan  personal  b r i e f o b s e r v a t i o n s on the  Green Mountain c o l o n i e s (numbers 3, 4, and 5, F i g 1) and Washington, Mt Heather, and  METHODS OF  I  observations  on  the  behaviour  April  The  until  21 September 1974.  vantage of  points,  between  50  without and  300  animals the m.  use  of 15-60  or 20-45 power.  and  from  were observed of  a  30 from  blind,  Observations  through b i n o c u l a r s of 7x35 or 10x40 m a g n i f i c a t i o n ,  least  Mt  of Vancouver  16 September 1973  scopes  on  B u t t l e r Peak (Fig 1, Table I ) .  I s l a n d marmots from 13 June u n t i l  distances  other  OBSERVATION  recorded  selected  1  or  at  were made spotting  Only o b s e r v a t i o n s o c c u r r i n g a t  15 minutes a f t e r my a r r i v a l  were i n c l u d e d i n my  results  —  a time I judged t o be s u f f i c i e n t f o r the marmots to h a b i t u a t e to my  presence.  preconstructed  Observations  were  recorded  t a l l y c h a r t s or with a tape  in  notebooks,  recorder.  on  11  Figure  1. L o c a t i o n s of known Vancouver I s l a n d marmot c o l o n i e s  1... Haley Lake s tudy area (1976)* 2. Green Mt, co lony 2 (1974) 3. Green Mt, co lony 3 (1974) 4. S k i C l u b , co lony 4 (1974) 5. S k i Club, co lony 5 (1974) 6. B u t t l e r Peak (197 4) 7. Jordan Meado ws (1930) 8. Mt Hhymper ( 1971) 9. Heather Mt ( 1974) 10. Shaw Creek, headwaters (1944) 11. Mt McQuillam (-Saunders; 1975) 12. Mt DeCosmos (?) 13. Mt M o r i a r i t y (1971)  14. Golden Eagle Basin (1910) 15. King Solomon Basin (1910) 16. Mt Douqlas (1910) 17. Mt Arrowsmith (1938) 18. Cameron Lake (?) 19. B e a u f o r t Range (1968) 20. Drink Mater Creek (1940) 21. Flower Ridge (?) 22. Golden Hinde (?) 23. Mt Albert-Edward (1970) 24. Mt S t r a t a (1955) 25. Mt Washington (1974) 26. Coraox (1968; a d i s p e r s i n g marmot was c o l l e c t e d near the c i t y )  * date p f the most r e c e n t c o n f i r m a t i o n o f colony e x i s t e n c e  11 a  32 km  12  Figure  2. Photographs o f the Haley Lake study area ft. Photograph of the whole Haley Lake c o l o n y B. Close-up photograph of the part o f the Haley Lake colony used most e x t e n s i v e l y by marmots {see a l s o F i g 22, 23, and 24)  12a  13  Table  I. Habitat c h a r a c t e r i s t i c s colonies  Colony Name land Number  North West L a t i t u d e Longitude  o f Vancouver I s l a n d marmot  Slope (Degrees)  Aspect  Elevation (Meters)  1 Haley Lake  490  01*  1240  19*  33-63<»>  SSE  1100- 1400  2 Green  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 S k i Club  49 0 03*  1240  20«  40<i>  E  1550- 1600  5 S k i Club  490  03»  1240  20*  3 5-56<* >  SSW  1550- 1600  6 B u t t l e r Peak  490  00«  1240  20*  35-70<2>  S  1500- 1550  7 Heather  49 0 53'  1240  30*  33<2>  S  1300- 1400  490  1250  14*  2 0< 2>  WSW  1500- 1600  Mt  Mt  8 Mt Washington  <f>  40 •  degree of s l o p e measured with a c l i n o m e t e r degree of s l o p e estimated  14  To  obtain  focal-animal sampling  quantitative  sampling  method  observed  at  1974).  Focal-animal  rarely  Often more than one  faced  with  (Ad l i b i t u m sampling, decisions  on  could  from  view  p e r i o d s were devoted  within  to scan  Altmann 1974).  when  o b s e r v a t i o n s on a f o c a l i n d i v i d u a l because disappeared  individual  one time, but i t was never p o s s i b l e t o observe  a l l animals a t a l l times was  (Altmann  c o n s i s t e d o f s e l e c t i n g one i n d i v i d u a l and c o n t i n u o u s l y  r e c o r d i n g i t s behaviour. be  data on marmot behaviour I used a  terminate  f o c a l animals u s u a l l y  a s h o r t time.  sampling;  to  I  the  Some o b s e r v a t i o n recording  of the  behaviour and l o c a t i o n o f each animal every 10 minutes.  ANALYSIS OF SOCIAL BEHAVIOUR DATA  I  c o n s i d e r e d a s o c i a l " i n t e r a c t i o n " t o be an u n i n t e r r u p t e d  s e r i e s of  social  individuals.  behaviour  Interactions  patterns were  or  "acts"  considered  a  social  behaviour  pattern  only  minute.  always  occurred  at  the  I  once unless i t was  separated by some other a c t other than t a i l r a i s i n g , s i n c e raising  two  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 o f more than one counted  between  tail  same time as some other a c t .  I n t e r a c t i o n s among t h r e e  animals  t r e a t e d as a s e t of dyads  ( i n t e r a c t i o n s between two a n i m a l s ) .  I  arbitrarily  interactions)  decided that a f o c a l animal sampling  had t o be g r e a t e r than 15 minutes include  (triadic  in  duration  before  were  period  I  i t i n e s t i m a t i n g the r a t e s o f i n t e r a c t i o n . , I f e l t  would that  15  by only c o n s i d e r i n g sampling periods that were minutes  I  would  eliminate  biasses  longer  inherent  than  i n short  15  samplinq  periods.  I c a l c u l a t e d the r a t e , R, that any group (i.e.  of  animals,  i ,  a d u l t males), performed any s o c i a l behaviour, B, from the  following  formula: Ri(B)  Si(B)  =  (1)  SHi  where Si(B)  i s t h e number of a l l o f t h e B a c t s i n v o l v i n q o r  performed by group when  i  individuals  during in  samplinq  qroup  i  periods  were  (S) ,  the  focal  i n d i v i d u a l s , and SHi i s the sampling  total  periods  time where  in  hours  (animal-hours)  i n d i v i d u a l s i n group  of  i were  the f o c a l i n d i v i d u a l s . The acts  r a t e , R i ( B ) , i s t h e r e f o r e an estimate that animals i n group  of  the  i were i n v o l v e d i n during  that they were a c t i v e above ground and i n p l a i n per  hour  t h a t I s a t observing  the c o l o n y ) .  are i n t e r a c t i o n s per animal-hour. individual  marmots  were  number  obtained  The r a t e s  view  of  B  each hour  (i.e.  not  The u n i t s of Ri (B) of  behaviour  by c o n s i d e r i n g  for  the qroup as  being composed of only one animal.  The  r a t e t h a t group  group j , i s :  i performed a c t B  with  any  specific  16  Si{B)i:j + Sj{B)i:j IR(B)i: j =  (2) (2)  (SHi + SH j)  where Si(B)i:j  is  between  the  i and j  participant  sura  of  during  from  all  B  a c t s that occurred  sampling  group  i was  periods  the f o c a l  when  the  individual,  . and t h e r e i s a 2 i n the denominator fact  that  by  watching  each  to  correct  group  e f f e c t i v e l y counted each i n t e r a c t i o n The  u n i t s of I R ( B ) i : j are i n t e r a c t i o n s  animal-hour,  where  groups  j were a c t i v e above ground and i n p l a i n view.  indicating  dyads  never  interacted  R i ( B ) i : j = 0,  even  of  thus  hours t h a t animals i n  during  sampling  though they may  during o b s e r v a t i o n p e r i o d s which were  of  a  periods,  have i n t e r a c t e d  shorter  duration,  i n d i c a t i n g t h a t R i ( B ) i : j i s g r e a t e r than z e r o .  estimate  I  twice.  represents  Some  number  the  independently  animal-hours i and  the  per  for  A non-zero  of R i ( B ) i : j can be c a l c u l a t e d i f S i ( B ) i : j i s c a l c u l a t e d  r a t h e r than counted  directly.  occurring  groups  between  i  The  total  and  j  number  over  all  of  B  acts  observations  r e g a r d l e s s of t h e i r d u r a t i o n i s T { B ) i : j , and the t o t a l number of B a c t s o c c u r r i n g between group  i and any  age  my  and  sex  is  T(B) i .  i n t e r a c t i o n s were unbiassed  If  other animal  observations  then S i ( B ) i : j  p r o p o r t i o n of S i (B) as T ( B ) i : j i s of T ( B ) i . Thus,  should  of  known  of b e h a v i o u r a l be  the  same  17  T(B) i : j  Si(B) i : j  T(B) i  Si(B) and  S i (B) Si(B)i:j  Substituting  =  x  T (B) i : j {3)  T{B)  eguation  i  (4) i n t o  S i <B)  equation  yields:  (3)  Sj(B)  x T (B) i : j  Equation  =  (4a)  (2) ( S H i + S H j )  is  4a  T(B) i : j  T(B) j  T (B) i IR(B) i : j  x  the  formula  I  used  to  calculate  interaction  rates.  It  was  necessary  interaction social  rates  behaviour  reciprocal.  I  reciprocal signals  if  behaviours  for  some  the  interactants  those  in  each  interaction  rate  estimate  j  was  the  since  same a s  the  Ri(G)i:j  social  the  which  other  rate  that  Ri(G) j : i  =  IR (G) j : i .  j  calculation patterns,  reciprocal  a  the  greeted  nonbe  exchange  of  see  acted  chasing).  Only  i  calculated greeted  with  also  Non-reciprocal  interactants  was  animal  since  to  greeting,  Patterns).  of  or  patterns  mutual  (e.g.  dyad  that  =  be  (i.e.  per  rate  behaviour  behaviour  to  Behaviour  i n the  either  appeared  from  act  be  act  Social were  explicit  social  could  differently  reciprocal  more  considered  the  on  be  patterns  between  section  to  i ,  one  for  with  very  each  animal  i.e:  and IB ( 6 ) 1 : However,  in  the  j  case  of  non-reciprocal  acts,  the  rate  that  18  animal at  i chased  which  animal  j chased  i ,  j i s not n e c e s s a r i l y  equal  to  the  rate  i . e :  R i ( C ) i : j * Ri(C) j : i . Thus  f o r  each  interaction  rate  non-reciprocal were  behaviour,  calculated  for  each  two  estimates  dyad,  IR(B)i:j  of and  IR(B)j:i. where Si{B)  x T(B)i:j  4.  Sj(B) x  T(B)i IR(B)i:j  T(B)i:j  T(B)i  = (2)  ( S H i + SHj)  and  Si{B)  x T(B) j : i T (B) i  IR(B) j : i  = (2)  The  interaction  get  the total  of  behaviour  interaction patterns  simple  relationship  summed  f o ra l l j's,  whereas  f o r a .specific dyad  rate  (i.e.  between  T(B)j • (SHi + SHj)  Ri(B)  because  c a n be added t o  or to get the rate a l l agonistic  i s  f o r any  acts).  and I R ( B ) i : j ,  IR(B)i:j  (4b)  a  when  group  There  i s no  IR(B)i:j i s  function  of  SHj  Ri(B) i s not.  The  formula  interaction by  rates  S j ( B ) x T(B) j : i  +  Armitage  rates  that yield  (1976a)  calculates  mean  combination  desired"  I  developed  t h e same r e s u l t s for  rates  have  some  dyads.  incorrectly  (Armitage  1976a).  by  for  as those  calculating  equations  However, adding  rates  used  Armitage i n "any  19  Unless otherwise  indicated, a significance  was used f o r a l l s t a t i s t i c a l  TRAPPING  0.05  AND MARKING  traps  Wisconsin, the  of  tests.  In 1974 marmots were captured live  level  manufactured 0. S, A.  i n 2 5x30x80 cm o r 22x22x63 cm  by Tomahawk L i v e Traps Co., Tomahawk,  B a i t s used were peanut b u t t e r , peanuts  and  l e a v e s and flowers of p r e f e r r e d s p e c i e s of food p l a n t s when  these were a v a i l a b l e (Appendix I ) .  Marmots were t r a n s f e r r e d from the t r a p s to a h a n d l i n g similar  to  the  one i l l u s t r a t e d by Taber and Cowan (1971).  t r a n g u i l i z a t i o n was necessary. flexible  steel  millimetre  p o s s i b l e i n an extended weighed  the  Animals were  tape  measured  while being  position.  I  occurrence  of  was a l s o recorded.  t h a t c o u l d be read  recorded  to  the  using  No a  held as n e a r l y a s the  marmot with a 12 Kg s p r i n g balance  Basle, Switzerland) The  cone  sex,  and  (Pesola S c a l e s , nearest  100  g.  e c t o p a r a s i t e s and a d e s c r i p t i o n of the molt Each animal was marked by a t t a c h i n g a s i n g l e  numbered r a b b i t ear tag ( s t y l e #4-1538) s u p p l i e d by the N a t i o n a l Band and Tag Co., Newport, Kentucky, U. S. A. or two c o l o u r e d Canada applied.  With each t a g one  p l a s t i c markers o f Dymo embossing tape  L t d . , Missisauga,  O n t a r i o ) , approximately  (Dymo  of  1x2 cm, were  Ear tags were p l a c e d as f a r from the margin o f the e a r  as p o s s i b l e before the t h i c k n e s s of  the  ear  became  limiting.  Tags were o c c a s i o n a l l y l o s t by other animals b i t i n g the c o l o u r e d markers  and r i p p i n g the whole tag from the e a r .  Tags were most  20  f r e q u e n t l y l o s t i n t h e t r a p s or d u r i n g h a n d l i n g .  Most  animals  were tagged at l e a s t twice d u r i n g t h i s study.  In  an  attempt  to  permanently  f r e e z e branding techniques  described  Winward  (1972)  (1966),  (unpublished). regrowth  of  recognized  Hadow  The d e s i r e d white  at  a  mark by and  result  of  animals I t r i e d t h e Farrell,  Roger,  Churchill freeze  Coburn  branding  is a  h a i r i n t h e shape of the brand which can be distance. ,  This  result  requires  melanocytes be destroyed but not t h e h a i r f o l l i c l e . laboratory  and  and  s t u d i e s , the regrowth  that  the  In previous  of white h a i r o c c u r r e d i n 3 t o  6 weeks and was r e t a i n e d through subseguent  molts  ( F a r r e l l et a l  1966, T a y l o r 1969, Hadow 1972, C h u r c h i l l and Coburn unpublished, Lazarus and Rowe 1975). a  1x5  cm  I used two brass branding " i r o n s : "  rectangle,  and  the other a c i r c l e , 3 cm i n o u t s i d e  diameter and 1.6 cm i n i n s i d e diameter. had  a  unique  surface  area  combination  of of  5 cm .  two  one  2  Each  brand  therefore  Each animal was marked with a  brands  o r r i e n t a t i o n and p o s i t i o n on the body.  by  varying  Brands  shaved area on the body f o r e x a c t l y 30 seconds.  the  were a p p l i e d  brand to a  21  MEASUREMENT OF MICROCLIMATE  In  1973  recorded.  In  obtained.  daily 1974  Daily  Gauge (Tru-Check record  of  maximum all  of  rainfall Inc.,  and the  temperature  following  was  Albert and  minimum  was  sere  measurements  were  recorded with a Tru-Check Lee  Minnesota).  humidity  was  u n t i l 3 November using a 31 day C a s e l l a thermohygrograph  temperatures  enclosed  A  continuous  obtained from  12 June  thermohygrograph.  i n a Stevenson  Screen  A d d i t i o n a l weather i n f o r m a t i o n  was  obtained  operated  for  the  Committee  the  of  instruments and  the were  Secretariat Province located  of  from  approximately  one.  Land  Columbia.  300  was  instruments  Environment  British  The  which  s i t u a t e d on a s m a l l eminence i n the c e n t r e of c o l o n y number  by  Rain  Use These  m below c o l o n i e s 5  6.  VOCALIZATIONS  I recorded v o c a l i z a t i o n s o f both trapped and marmots with a Uher 4000IC Report used  was  either  an  ElectroVoice  microphone or a Sony Dynamic speed was  always 19.05  recordings both  on  trapped  calculations were from  cm  a  Kay Co.  and  free  of  tape r e c o r d e r .  whistle  M136  per s.  64 4  The  Sound Spot  microphone.  ranging  microphone directional  The  recording  Sonograms were made from  M i s s i l y z e r model 675. ranging  free  marmots  were  l e n g t h and freguency.  f r e e ranging marmots o n l y .  these  Recordings used  in  of the  A l l other data  22  RESULTS  HABITAT CHARACTERISTICS  Vancouver I s l a n d a a r i o t s l i v e i n are  characterized  by  steep  subalpine  cliffs,  talus  meadows t h a t are u s u a l l y o r i e n t e d south of (Table I , F i g 2 and steep,  the  3).  the  and  Where  snow  slope  is  and  open line  becomes l e s s  herbaceous p l a n t steep  enough,  creep i n h i b i t the establishment of t r e e s .  Evidence of these f o r c e s can and  the  that  east-west  Below the t a l u s , the slope  develop.  avalanches  saplings  debris,  s u b s t r a t e becomes more s t a b l e , and  communities  habitats  be seen i n  the  form  of  uprooted  the d i s t i n c t b a s a l crook i n a l l e s t a b l i s h e d  trees  (Fig 3A) . ,  A l l of  the  communites  that  marmot  colonies  that  I  visited  had  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 o f B r i t i s h Columbia Peterson, and  K r a j i n a 1970).  Within  et a l (1970) d e s c r i b e e i g h t p l a n t the Haley  Lake  study  fflgrtengiaS5'  yellow  huckleberry  Vaccinium  Bgctinata, Appendix  and I)  mertensianae mertensianae.  plant  area  as  mountain  P l a n t s found hemlock  cedar Chamaecyparis n o o t k a t e n s i s . delicjosum,  mountain were  the Parkland Subzone Brooke  associations.  such  daisy  of  peregrinus the  subassociation  colonies.  Tsuga  Leutkea (see  also  Nano-tsugetum nano-tsugetum  Other plant a s s o c i a t i o n s were a l s o present  Haley Lake study area and on other  on  blue-leaf  partridgefoot  Eriaeron  characteristic  association,  (Brooke,  on  Three c o l o n i e s ,  the Mt  23  Figure  3. Photographs o f Vancouver I s l a n d marmot h a b i t a t 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 h a b i t a t B. The Mt Washington colony illustrating the dense cover of white rhododendron and a l p i n e f i r  23a  B  24  Washington, Heather Mt> and C e n t r a l Green Mt (colony 3) d i f f e r e d from  the others i n  meadows  were  Washington. area  they  were  characteristic  of  not  below  The area was a l s o covered  rhododendron  Rhododendron  by  a  a l p i n e f i r Abies dense  growth  of  a l b i f l p p u m . and Vaccinium sp.  3B) .  The: c l i m a t e o f characterized  by  the  cool  Subalpine short  length  of  the  growing  Mountain  summers  considerable p r e c i p i t a t i o n f a l l i n g season  as is  and  Hemlock wet  snow  difficult  above  f r e e z i n g while snow s t i l l  great accumulations Table  is with  II).  The  to determine i n are  often  well  p e r s i s t s on the s i t e due t o the  d u r i n g the winter  (Brooke  et a l  1970  and  I I ) . On the Haley Lake study area i n 1974, the number o f  f r o s t - f r e e days was above average the e f f e c t i v e growing season the  Zone  winters  (Table  s u b a l p i n e areas s i n c e minimum a i r temperatures  colony  (Table  II)  at 135 days  (Table  above  the  colony  were  as  a r e s u l t of a p e r s i s t e n t snowpack.  too  The  cliffs  The  than  the  rest  the  the  A s m a l l part of colony 2 was f r e e o f  snow very e a r l y i n 1974, a p p a r e n t l y having been kept by  of  on t h e other c o l o n i e s were a l s o the f i r s t  areas to become snow f r e e .  free  cliffs  s t e e p to accumulate much snow and  t h e r e f o r e became snow f r e e much e a l i e r colony.  I I ) , but  was about two weeks l e s s on much o f  However, the snow pack was not evenly d i s t r i b u t e d .  snow  Open  a l l marmot c o l o n i e s except Mt  with numerous s m a l l , 1-5 m, t r e e s , mainly  white  cliffs.  Marmot burrows on Mt Washington were l o c a t e d i n an  lasiocarpa.  (Fig  that  prevailing  wind.  relatively  Marmots foraged on these  e a r l y snow f r e e areas u n t i l the snow melted  from  the  main  part  25  Table  I I . C l i m a t i c c h a r a c t e r i s t i c s o f the Haley Lake study area and a t y p i c a l Parkland Subzone l o c a t i o n i n the Coast Mountains, a f t e r Brooke e t a l 1970  , Climatic Characteristics  Baley Lake Study Area  Year  Brooke e t a l 1970  285 35%  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)  25  1974  50  mean temperature J u l y through September (°C)  13.0 15.5  1973 1974  12.0  13-20 June 3 June  1973 1974  end o f May  date of the l a s t f r o s t i n the spring number o f f r o s t - f r e e days  115 135  1973 1974  114  maximum accumulation of snow (cm)  approximately 300  1974  370  1973 1974  l a t e June  l a s t accumulation o f show i n the s p r i n g  e a r l y June l a t e June  26  of the c o l o n y .  On the Haley Lake study area i n 1974  to  the  forage  on  cliffs  for  marmots had  6 weeks a f t e r emergence i n the  spring.  Steep s l o p e s habitat  are c h a r a c t e r i s t i c of Vancouver I s l a n d  because they are s u s c e p t i b l e t o avalanches.  provide  suitable  communities reducing  habitat  through  the  1) by  Avalanches  maintaining  inhibition  marmot  herbaceous  of t r e e growth, and  the accumulation of snow, r e l a t i v e  to  2)  flatter  areas,  which  in  in the  s p r i n g , thus i n c r e a s i n g the e f f e c t i v e growing season.  The  by  t u r n r e s u l t s i n the meadow becoming snow f r e e e a r l i e r  average  area  h e c t a r e s (range 0.5  to  of  8  marmot  colonies  was  about  two  4.0).  PHYSICAL CHARACTERISTICS  I  recognized  marmots:  infants,  Identification  of  did not t r a p any easily  the f o l l o w i n g age yearlings, age  distinguished  sample s i z e was  because  females  female  study,  adults.  weights (Fig 4 ) . but  infants  of t h e i r small size  female y e a r l i n g s , but  weighed l e s s than two-year-olds  than  this  based on  and  I  were  (Fig 4).  My  too s m a l l to compare t h e d i f f e r e n c e s between the  weights of male and  Some  two-year-olds,  c l a s s e s was  i n f a n t s during  c l a s s e s of Vancouver I s l a n d  had  yearlings  y e a r l i n g s as  (F=123.13; d.f. = 1,16;  a  p<0.001).  weights t h a t were s i g n i f i c a n t l y (F=65.6;  d.f.-1,13;  group  greater  p<0.001)  but  27  Figure  4. Seasonal changes I s l a n d marmots  in  the  weights  of  Vancouver  The growth r a t e equations a r e as f o l l o w s : A d u l t s , sexes combined  y = 0.0245x • 2.466  Two-year-old females  y = 0.0259x + 1.516  Yearlings,  y = 0.0230x + 0.584  sexes combined  where y i s t h e weight i n kilograms and x i s t h e number o f days s i n c e (30 A p r i l )  spring  emergence  * known aged two-year-old i n f a n t female; data from a specimen c o l l e c t e d from Mt Washington i n 1965, U n i v e r s i t y of A l a s k a specimen #28754 2  2?a  28  significantly P<0.001). 1975,  l e s s than the l a r g e s t females  I assumed that these animals were two-year-olds.  I  captured  weighed s l i g h t l y from the'growth consistent  M. c a l i g a t a  absence  one  r a t e equation the  Two-year-olds  adult  (Fig  4).  This  predicted  observation  is  that two-year-olds weigh l e s s  can  be  recognized  (Barash 1974b) and M. olympus  by  weight  in  (Barash 1973b).  The  of any males r e c o g n i z a b l e as two-year-olds c o u l d be the  the  of any  y e a r - o l d males are s t i l l of  two-year-old  males  on  weight  as two-year-olds.  In  o l d males on my study a r e a s .  to  Adult males weighed s i g n i f i c a n t l y  Growth  rate  then,  two-yearrefers  to  be at l e a s t t h r e e years o l d . more than a d u l t  females  (Fig  p=0.014).  appears  c l a s s e s throughout the covariance indicated  to  summer  (1973b)  be  linear  (Fig  4  and  that the r a t e of weight  s i g n i f i c a n t l y among sex and age  Barash  two-  I assumed t h a t the same would  The term a d u l t ,  considered  5; F=6.87 d.f. = 1,28  their  d i s t i n g u i s h a b l e from a d u l t males on the  (Barash 1973b).  are  study  M. olympus*  be t r u e f o r Vancouver I s l a n d marmots and that I had no  that  my  growth of males being such t h a t they reach  (heaviest) weight  animals  She  males t h a t were n o n - y e a r l i n g s had s i m i l a r weights.  or  basis  In  known to be two years o l d .  assumption  r e s u l t of the absence areas  female  l e s s than the average two-year-weight  with  than a d u l t s .  All  (F=18.7; d.f.=1,22;  for 5).  a l l sex and  age  Analysis  of  gain d i d not  differ  classes.  used the change i n M. olympus- tooth c o l o u r  29  Figure  5. Seasonal changes i n the weights of a d u l t marmots The  growth r a t e equations are as f o l l o w s :  adult  males  y = 0. 0210x «• 2. 56 1  adult females y = 0.0263x • 2.489 where y i s the weight i n kilograms and x i s t h e number o f days s i n c e s p r i n g emergence {30 A p r i l ) .  29a  3 V  B-1  7-1  • A Y S AFTER SPRING EKERGEISCE  C30 A P R I L )  30  (from  dull  classes. since  the  white  t o dark  This  was  tooth  orange)  not  as  an  possible  colour of  aid  in  separating  i n Vancouver  a l l Vancouver  Island  Island  marmots  age  marmots  was  dull  very  dark  except  for  white.  The  fresh  brown. a  of  M.  vancouverensis  This colour i s uniform  conspicuous  small the  pelage  white breast  gradually cinnamon  patch  of  white  mark  on  the  and  fur  over  brown.  the  The  of  the  and  and  Island  body  nose  and  white  of  the  following  marmots  or  the  some  colour  summer  Vancouver  most  around  forehead,  abdomen.  fades  over  i s black  mouth,  a  streaking  on  fresh  spring  molt  pelage  to  only  a  light  once  per  year.  The field old  because faded  July. and  progression  The  the  body  usually animals  hibernated this  time.  the  next  previous July. previous  on  rump,  on  The  emergence  extremely of the  and  to  the  at  the  no  subsequent  molt  last  tail,  and  animals  same s t a g e  had  began  their  nonetheless,  molted  of  change not  first  or  pelage  The  complete  September;  individual the  fresh  variable.  head,  appear  of  the  over  the to  the  molt  by  in  the  the the mid-  forelegs rest molt  rump.  molting  of were Some  time  finished  they at  emerged  from  hibernation!  molt  they  were  as  occurred  completed in  the  areas  in  with  first  Incompletely  year,  sharply  f u r emerged  in late  an  c o n t r a s t e d so  observed  molting  not  fall,  easily  signs of  spring  If  pelage  was  showed  back  did  molt  first  was  the  the  Animals  shoulders.  the  fresh  fur.. new  of  the  until  i t s molt areas  at  the from  having  the next the the  31  oldest  pelage.  black f u r . summer  Young  animals  They d i d not  but  appear  sex  to  in  I  from the burrows with  molt  molting would be d i f f i c u l t  absence o f any c o l o u r change. differences  emerged  their  first  t o d e t e c t because  o f the  could  during  not  detect  any  other  molting c h a r a c t e r i s t i c s among d i f f e r e n t age and  classes.  Davis the  (1966) noted t h a t M. mpnax a l s o had a v a r i a b i l i t y  progression  rump.  of  the  molt, although i t always began on t h e  He a l s o observed that some animals d i d not complete  molt each year.  One molt per year appears to be the r u l e  marmots, W. raonax (Hamilton 1934, Davis 1966), (this  study),  (Walker  M. f l a v i v e n t r i s  among  (Armitage 1974), and M. olympus (1973b) f o r  f r e e z e brands were a p p l i e d on e i t h e r the 23, 24, or 25  J u l y 1974.  T h i s was the e a r l i e s t date p o s s i b l e because  a labour  s t r i k e prevented the p r o d u c t i o n of dry i c e u n t i l t h i s time. the  At  time of a p p l i c a t i o n , f r e s h pelage had already begun t o grow  over much of the body. fur  their  vancouverensis  1964) , although t h i s was disputed by Barash  All  in  had o c c u r r e d .  Four weeks a f t e r branding no regrowth o f  The branded  areas were bare, the  and a l a y e r o f s k i n having been sloughed o f f . branding, the brands had s t i l l which  was  in  the  process of growing  with white t i p s .  white f u r i n 1974.  In  hairs  Seven weeks a f t e r  not grown i n completely.  The f u r  i n was e i t h e r b l a c k , t h e  normal c o l o u r o f f r e s h pelage, white, a s black  club  expected,  or  rarely,  No brands were v i s i b l e at a d i s t a n c e as June  1975  the  brands  had  still  not  32  regrown  on  the  were p o s s i b l e . were  one i n d i v i d u a l f o r which d e t a i l e d o b s e r v a t i o n s No animals d i s p l a y e d white brand markings  v i s i b l e from a d i s t a n c e at t h i s time.  brands on  one  individual  pigmentation.  However,  had three  In August  regrown  with  marmots  had  1975,  normal  power  s p o t t i n g scope.  the  coloured  brands  that were  i d e n t i f i a b l e at a d i s t a n c e o f about 100 m when viewed 15-60  which  through  a  On c l o s e o b s e r v a t i o n I found that  these brands were only s p a r s e l y covered with white f u r .  Since on Vancouver  I s l a n d 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 later,  freeze  t h i s study.  branding  However, i n  was u s e l e s s as a marking the  future  the  branding might be i n c r e a s e d i n two ways.  technigue f o r  efficacy One,  of  freeze  branding c o u l d be  done e a r l i e r i n the summer t o minimize the time between branding and  the  beginning  of  the  molt.  becoming v i s i b l e  i n t h r e e to s i x  studies  1972,  (Hadow  and Rowe 1975). length  of  time  very  which  Hadow  Free ranging  brand to  in  the  sguirrels  (Sciurus a b e r t i ) and  was  laboratory  applied.  achieve  1972,  marmots  time  1973)  the  important  branding  (Hadow  as  C h u r c h i l l and Coburn u n p u b l i s h e d , Lazarus  during  et a l 1966,  unpublished).  weeks  i n brands  A second improvement t h a t c o u l d be made i s  a p p l i c a t i o n time i s (Farrell  T h i s could r e s u l t  good  Churchill may  reguire  The  results  and a  the  Coburn  different  than was p r e d i c t e d from experiments with Abort*s  and  Columbian  fox  squirrels  ground  (Sciurus  sguirrels  £2l]iSfei§llSS; C h u r c h i l l and Coburn unpublished) . from my r e s u l t s whether the branding time t h a t I  niger)  {Spermophilus I t i s not c l e a r used  was  too  33  short  or  too  long  ( i f either).  S i n c e the brands took a long  time t o regrow and normal  pigmentation sometimes developed,  a p p l i c a t i o n time appeared  to be too l o n g (Hadow 1972).  the  resulting  intermediate  pigmentation  in  the  However,  some  animals  i n d i c a t e d that the a p p l i c a t i o n time was too s h o r t ( C h u r c h i l l and Coburn u n p u b l i s h e d ) .  Lazarus and pressurized  Bowe  (1975)  refrigerant  as  used a  a  freeze  commercially branding  available  agent.  Their  r e 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 f i e l d conditions. similar  dry  ice,  especially  under  A p r e s s u r i z e d r e f r i g e r a n t a v a i l a b l e i n Canada  t o the one used  Refrigerant  using  by Lazarus and Bowe (197 5)  i s Can.O.Gas  12, V i r g i n i a Chemicals I n c . , Portsmouth,  Va.  U.S.A.  COLONY COMPOSITION  I c o n s i d e r e d a c o l o n y t o be a group isolated  both  localities. colonies are  of  one  I was unable  the  Table I I I ) . be  and two  socially  on Green Mountain  from  (Fig 1).  Since I d i d  not  that  was  other  such  colonies,  These r e s u l t s  trap  animals  in  only able t o r e c o g n i z e i n f a n t s and o l d e r animals and t o determine t h e i r sex.  most animals i n 1974.  before  and  animals  I knew the exact p o p u l a t i o n s i z e f o r two  presented i n Table I I I .  1973, was  geographically  of  The  However, I knew the s t a t u s  average  colony  i n f a n t s emerged above ground,  was  size 8.3  in  June,  animals  A l l the other c o l o n i e s t h a t I observed appeared  of a s i 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  (n=4. to 3.0  34  ±.32  (SE)  infants  (n=5) .  Movement of marmots between c o l o n i e s was were  only  four  occasions,  the  way  from  and two  t r a c k s of marmots  on 20 May  1974  in  and most of  colony one to colony three on 11 June 1974.  adult male was  observed  June  seen  he  There  t h a t I knew of, when marmots moved  between c o l o n i e s on Green Mountain., I saw the snow between c o l o n i e s one  very r a r e . ,  was  on colony one  on  colony  two  u n t i l 2 June 1974. where  One  On  12  he remained f o r the  d u r a t i o n of the summer., An a d u l t male immigrated t o colony from  an  unknown  l o c a t i o n on, or s h o r t l y before, 25 June  He remained t h e r e f o r the d u r a t i o n of the therefore  was  occupied  when and  I was  unable  Colony  two  present  at  any  never  one  one  time  t o determine the sex of three marmots  I f i r s t captured them., They were r a r e l y seen were  1974.  by t h r e e d i f f e r e n t a d u l t males i n June  197,4; however, there were only (Table I I I ) .  summer.  one  recaptured;  therefore,  their  subsequently  sex  was  never  determined. ,  ACTIVITY PATTERNS  The guite  g e n e r a l p a t t e r n of Vancouver I s l a n d marmot a c t i v i t y  similar  (Armitage Island  1962,  to  Gray 1967,  marmots  activity  have  an  reported Barash annual  for  other  marmot  1973b, Hayes 1976). schedule  species Vancouver  made up of a summer  period of 4 t o 5 months and a winter h i b e r n a t i o n p e r i o d  of 7 "to 8 months. first  that  is  day  In 1974  that I v i s i t e d  marmots were a c t i v e on 30 A p r i l , the Haley Lake study a r e a .  the  Tracks i n  35  Table  I I I . Aqe  and  sex  composition  Number Cclcny  Year  Month  1973  June July Auq Sept.  1974  1973  197 4  Hay June July Aug Sept  AM  AP  A = Adult M = Male  of A?  marmot c o l o n i e s  Marmots 2F  2?  cf  one  2 2 2 2 2  2 2 2 2 2  1 1 1 1  Aqe  and  YF  Y?  I?  YH  6 4 4 1 1 1  1 1  1 1 1  1  2 2 2 2 1  6 4 4  1 1 1  1 1 1 1  1 1  ? = Sex unknown 2 = Two-year-old  ard  Each  11 11 8 8  June July Aug May June July Aug  of  1 1 1  1 1 1 1  I = Infant • Y = Yearling  Sex TOTAL  11 17 12 12 11 11 10 8 6  2 2 1 1 1 1  two  6 6 6 4 5 4 4  1 -  Female  36  the snow i n d i c a t e d t h a t  marmots had probably not been a c t i v e f o r  very long and a l l i n d i v i d u a l s had not hibernation had  at  this  time.  necessarily  ended  There was no sign t h a t any  their animals  yet emerged when I v i s i t e d the Haley Lake study area  April  1975.  I  September 1974. several  ;days  last  saw  Thus  1972  I  17  marmots on 16 September 1973 and  I observed v a r i o u s c o l o n i e s f o r many hours, after  these  dates.  marmots on the Haley Lake study October  on  early  area  May  on  N a t u r a l i s t s recorded s e e i n g on  30  September  (Bob M o r r i s and Ted Barsby p e r s o n a l  considered  21  to  and  1  communication).  be a r e a s o n a b l e estimate of  s p r i n g emergence with most i n d i v i d u a l s i n h i b e r n a t i o n  again  by  mid-September.  I  did  not  notice  d i f f e r e n c e s among age-sex c l a s s e s with  r e s p e c t t o the time of s p r i n g emergence or f a l l I  have  few o b s e r v a t i o n s from these p e r i o d s .  unusual i f Vancouver I s l a n d marmots d i d not emergence all  I t would be  have  age  but guite  specific  and h i b e r n a t i o n times s i n c e t h i s i s c h a r a c t e r i s t i c of  other marmot and ground  (e.g.  Vos  1972,  Yeaton  were  born  and G i l l e s p i e 1972, on  the  sguirrel  1960,  Barash Haley  1962,  that  I  know  of  Iverson and Turner  Both l i t t e r s  of i n f a n t s  that  Lake study area i n 1973 emerged from time  t h a t i n the Mt Douglas  t h i r d week i n J u l y  species  Armitage  1973b).  t h e i r burrows f o r the f i r s t noted  hibernation  on  11  July.  area no l i t t e r s  Swarth  (1912)  had emerged by the  1911.  I compiled a c t i v i t y budgets  f o r each i n d i v i d u a l i n terms of  the percent of the t o t a l o b s e r v a t i o n time that  a  marmot  spent  37  resting the  (lying  burrow,  behaviour,  s i t t i n g o u t s i d e of the burrow), f e e d i n g , i n  moving and  months o f May 10.  or  (without  grass  feeding),  collecting.  engaging  Activity  in  social  budgets  f o r the  through September a r e presented i n  Figures  to  Only t h e data f o r r e s t i n g , f e e d i n g , and i n burrow time are  presented s i n c e these t h r e e behaviours accounted f o r 93%  more  than  of the time budgets of a l l animals i n each month.. The data  were  combined  significant  for  a l l individuals  differences  these t h r e e a c t i v i t y  among  since  no  with the time of year. May  and  September.  J u l y , and August marmots were much more a c t i v e i n the  mornings and evenings than at midday. midday  were  patterns.  A midday a c t i v i t y l u l l was not apparent i n June,  there  age-sex c l a s s e s with r e s p e c t to  The d a i l y p a t t e r n o f a c t i v i t y v a r i e d  In  6  activity  in  The  obvious  decline  in  J u l y and August seemed t o be the r e s u l t of  three f a c t o r s ; temperature, an i n h e r e n t c i r c a d i a n rhythm, and general  decline  in  the  amount  of  a  time spent f e e d i n g as the  summer progressed.  ....... I d i v i d e d the day i n t o t h r e e p e r i o d s , morning, evening,  based  on the d u r a t i o n of a marmot day.  midday,  These p e r i o d s  v a r i e d s l i g h t l y between months but the midday p e r i o d was between 1100 and  1600  hours.  The  maximum  daily  and  usually  temperature  occurred during the midday p e r i o d except on a few u n u s u a l l y c o o l days.  The  percent of time spent feeding at midday dropped o f f  d r a m a t i c a l l y when the maximum d a i l y temperature exceeded  20°C.  38  Maximum D a i l y Temperature (°C) 0 15 20 25  -  14.9 19.9 24.9 29.9  23 22 6 0  a l s o , the time spent correlated  of Time Spent Feeding the Midday P e r i o d  Percent During  with  in  the  burrow  at  midday  the maximum d a i l y temperature  was  directly  {r=0.69, p<0.01).  Temperatures over 20°C were much more common i n J u l y and than i n the other  When still the  the  months.  maximum  temperature  was  l e s s than 20°C marmots  fed sigi.fica.ntly l e s s d u r i n g midday than they morning  and  p e r i o d s were t=2.90, daily  evening  lumped  df=63, f ee di ng  august  periods  and  during  {data from morning and  evening  tested  against  Thus  there was  p<0.01). activity  did  which  was  the  midday  period;  a bimodal p a t t e r n of  accentuated  by  high  temperatures.  The  total  the summer. in  both  time spent  T h i s trend was  food  guality  summer progressed melted  and  decreased,  the r e s u l t of  throughout  an  increase  guantity  over the summer.  vegetation  guantity  increased  guality  as  As  the  and  are  selected  by  as the time r e q u i r e d t o there  would  further accentuatinq  be  less  the snow  increased as more s p e c i e s came  Flowers are more n u t r i t i o u s than v e g e t a t i v e  1972)  {Appendix I ) .  probably  decreased  and  vegetation  i n t o flower. (Svoboda  fe e d i n g per day  marmots obtain  pressure  parts  when a v a i l a b l e sufficient  food  to feed at midday,  the a c t i v i t y l u l l s i n J u l y and  august.  The  39  Figure  6. A c t i v i t y budgets f o r May, 65 animal-hours of  a l l animals combined  observation  39a MAY ACTIVITY CALL ANIMALS) 100. 90-.. 80... 70... 60...  50-. 40... 30-..  ao... 10...  0.  10-  11 > 1H>  13-  14-  IS-  ! £ • 17-  18-  19. BO-  17-  IB-  IS-  17.  IS.  19- 5 0 .  HOUR DF THE DAY 10O90-.. 80... 70-.. GO...  50v. 40-.. 30...  30'  10 0  10.  i l .  12-  13.  14.  15.  IB-  BO  HOUR OF THE DAY 100. 90. 8070GO50-, 40-.  30-. 50.. 10-. 0..  10-  11-  12-  13-  H  14.  X  IS.  R OF THE DAY  40  Figure  7.  Activity 175  budgets  f o r June, a l l animals  animal-hours of  observation  combined  40a JLNE ACTIVITY  CALL ANIMALS)  100. 90... BO... 70... 60... 50... 40... 30-.. 30... 10... 0-._ 4-  10-  IS-  11.  13.  14-  I-CUR CF THE  15-  16-  17.  16>  17.  IB'  19-  SO-  HI.  33.  SS-  19-  50.  51-  HE-  S3-  DAY  10090. 8070-  eo504030.  so. 10.  o.  10.  11.  15 < 13-  14.  HOUR OF THE  15.  18.  OAY  100-. 90-. 80-. 70-. 60-. 50-. 40". SOSO-. 100-  9.  10-  11-  IS"  13-  14-  15>  HOUR DF THE DAY  1G-  IB-  19.  SO-  El.  E2«  S3>  41  Figure  8. A c t i v i t y budgets  f o r J u l y , a l l animals combined  202 animal-hours of  observation  4la JULY ACTIVITY  CALL ANIMALS)  10090... BO... 70:. GO... 50... AO:. 30... 50... 1X3... 0.._ 9-  10.  ! ! • 15-  13-  14-  15-  16-  17-  IS'  19-  SO-  5 1 « SH- B3>  HOUR OF THE DAY 10090-.. 90-.. 70-.. 60... 50... 40«.. 30-.. 50-.. 10-.. 0...  6*  7.  6>  10.  1i l ' '1 l S . 113 •  14.  15-  IB-  17. IB'  19-  SO-  51 • 5 5 ' S3-  HOUR OF THE DAY 10090'.. BO-.. 70... GO 50... 40... 30... SO-.. 100 4.  5-  10.  11.  12-  13-  14-  15-  HOUR CF THE DAY  IB-  17-  IB'  19-  50-  51-  25- S3-  42  Figure  9. A c t i v i t y budgets f o r August, a l l animals combined 114 animal-hours o f o b s e r v a t i o n  42a AIGLET ACTIVITY  CALL ANIMALS)  100. 30-.. 80-.. 70... BO. . 50... 40. . 30-.. SO... IX)... 0-._ 10.  11.  IS-  13-  14-  HOJR OF  15-  IB-  IB-  13-  SO-  51-  SB-  53-  THE OAY  10090... 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 10090SO7060-. 504030HO100-. 10.  11.  IH.  13-  14-  15>  HXR CF THE DAY  IE-  17-  18-  19-  HO-  HI.  EE-  S3-  43  Figure  10.  Activity 70  budgets  f o r September,  animal-hours of  observation  a l l animals  combined  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-  16-  17.  IB-  19.  20.  21.E2  22  HOUR OF THE DAY  100 90-.. SO... 70-.. BO-..  50 40-.. 30... 20... 10... 0  10.  11.  12-  13-  14.  IS.  HOUR OF THE DAY 10090-.. BO... 70...  BO. 50... 40... 30... 20... 10...  0. R-  10-  11-  12-  13-  14.  15-  HOUR OF THE DAY  IB-  17.  IB-  13-  BO.  21.  22.  44  absence of a midday a c t i v i t y l u l l of s h o r t day  l e n g t h or metabolic  i n September may  be an  effect  changes of upcoming h i b e r n a t i o n  on the b a s i c c i r c a d i a n rhythm.  I and I saw  saw  marmots  c o l l e c t i n g the brown dead stems of  sedges and t a k i n g them i n t o t h e i r burrows on 152 assumed t h a t t h i s m a t e r i a l was marmots e a t i n g i t above  collecting  decreased  ground.  The  frequency  grass  at a l l times of day,  most o f t e n a t the very end of the day, the burrow f o r the  of  grass  a  slight  through the summer but there was Adult  c o l l e c t e d more o f t e n than the other age-sex c l a s s e s . collecting  occasions.  used f o r bedding s i n c e I never  i n c r e a s e i n September j u s t p r i o r to h i b e r n a t i o n .  marmots  grasses  females  I observed  but i t occurred  j u s t b e f o r e they  entered  night.  VOCALIZATIONS  Whistles  The  sound  most  frequently  produced  marmots i s a loud p i e r c i n g " w h i s t l e " vocal  chords.  The  (n=36).  about  5700  these  harmonics  fundamental  Hz  and  which  originates  in  dominant frequency of a w h i s t l e i s the  harmonic or fundamental frequency. 2 9 1 0 ± 3 3 Hz  by Vancouver I s l a n d  Whistles a third  contain  frequency  The  first  the first  harmonic occurs  a l s o possess a second harmonic harmonic at about 8400 Hz.  much  (Fig  sound d i d not vary a p p r e c i a b l y  less  11  and  energy 12).  The  throughout the  than  at  Both o f  does  intensity of duration  at  of  the the the  45  Figure  11. Representative  sonogram of a short  Recording speed: 19.05 cm per s Playback speed: 4.76 cm per s Narrow bandwidth f i l t e r  whistle  eoooo. 1BOOO-. 1GOOO-. 14O0O-.  iaooo._ 10000..  TIME  (SETONDS)  46  Figure  12. R e p r e s e n t a t i v e  sonogram o f a l o n g  R e c o r d i n g s p e e d : 1 9 . 0 5 cm p e r s P l a y b a c k s p e e d : 9 . 3 5 cm p e r s Wide b a n d w i d t h f i l t e r  whistle  46a  47  whistle.  I subjectively classified in  the  field  duration.  as  being  e i t h e r " l o n g " , "medium", or " s h o r t " i n  I then used sonograms t o determine  each of these w h i s t l e types. whistle classification  mean duration  long medium short  The  The  (s)  to  distinguish  long  or  short.  duration  of  r e s u l t s were:  0.27 0.23 0.12  s i n c e most w h i s t l e s t h a t I  the  range (s)  0.57 0.26 0.20  inability  either  most marmot w h i s t l e s t h a t I heard  - 0.84 - 0.29 - 0.26  SE  n  0.052 0.009 0.009  14 7 17  medium w h i s t l e s was  heard  in  the  field  not  serious  were  clearly  Medium w h i s t l e s were t h e r e f o r e  omitted  from subsequent a n a l y s i s except where noted.  There was trapped  and  (t= 1. 50)  no s i g n i f i c a n t  free  ranging  or freguency  ft w h i s t l i n g  d i f f e r e n c e between the w h i s t l e s of  marmots  with  respect  sequence  was  considered  to be any minute  sequence of long w h i s t l e s was  400  whistles.  The  the l e n g t h of the w h i s t l e . w h i s t l e i n t e r v a l of 17.9  Most  (Fig 13).  and  contained  i n t e r v a l between w h i s t l e s v a r i e d Long  ±1.59  whistles  s and  The  50 w h i s t l e s i n 9 minutes.  sequence of short w h i s t l e s l a s t e d 28 minutes  about  w h i s t l e or  apart.  w h i s t l i n g sequences c o n s i s t e d of a s i n g l e w h i s t l e  One  duration  (t=l.05).  group of w h i s t l e s t h a t were l e s s than one  longest  to  had  a  mean  with  inter-  short w h i s t l e s averaged  2.9  48  Figure  13.  ft comparison of the w h i s t l i n g sequences  length  of  long  and  short  BO.  1  2  NUMEER C F WHISTLES  3  4  5  >5  I N A WHISTLING SEQLENCE  4.9  ±0.08 s between w h i s t l e s .  Marmot  whistles  serve  (Armitage 1962, Waring 1966, Island black  marmots bears  (irsus  chrysaetos), t a i l e d hawks are  always  bald  the  On one o c c a s i o n on a marmot. a  warning 1975)..  Vancouver  (Felis concolor),  golden  (Haliaetus  function  eagles  (Aquila  leucocephalus),  were d e t e c t e d .  or r e d -  These  species  major p r e d a t o r s o f Vancouver I s l a n d marmots.  On another, at  A l l of these  other  marmot s p e c i e s 1974;  1973b,  americanus)  I saw a golden  marmot  a  w h i s t l e d when cougars  eagles  view.  Banfield  Barash  (Buteo j a m a i c e n s i s )  probably  caught  primarily  I  a  eagle make an u n s u c c e s s f u l  am  guite  that  two  cougars  spot where t r e e s p a r t i a l l y obscured  predators have (cougars,  golden  sure  attack  eagles,  been  reported  to  prey  Barash 1973b, 1975; black Olendorff  1976,  my on  bears,  Barash  1975,  Armitage and Downhower 1974; b a l d e a g l e s , Beebe 1974; r e d - t a i l e d hawks.  Bent  1937)..  species,  coyotes  arctos).  are  Two  (Canis  absent  important latrans)  from  predators of other marmot and  grizzly  Vancouver i s l a n d  fOrsus  bear  (Cowan and Guiguet  1965) .  Whistles were given only 27 of 61 times (56%) t h a t raptors  (Faloniformes)  and  ravens  (Corvus corax) were present  (Table IV) .  whistles  also  occurred  causes,  but  I  not  associate  whistles  (Table IV) .  could  smaller  occasionally  for  other  any cause f o r 59% o f the  ;  Since such a l a r g e p r o p o r t i o n of  whistling  seguences  was  50  Table  IV. Causes of whistling and keeaw sequences  Stimuli  Whistling Sequences Number Percent  Potential Predators black bears ccugars l a i d eaqles qclden eagles unidentified eagles r e d - t a i l e d hawks subtotal  13 3 4 5 6 5  Snail Raptors and Havens marsh hawks sharp-shinned hawks Cooper's hawks sparrow hawks unidentified small raptors ravens subtotal  10 4 3 1 4  Miscellaneous Causes i n t r a s p e c i f i c chases ai r c r a f t b l a c k - t a i l e d deer people l a n d - t a i l e d pigecns? ccmmon f l i c k e r s ? Canada jays? snow and rock s l i d e s sudden h a i l storm subtotal  36  5  27 5 4 4 3 1 2 1 2 1  23  6.1 1.1 1.9 2.4 2.8 2.4  Keeaw Sequences Number Percent  17.0  4.7 1.9 1.4 0. 5 1.9 2. 4  2.4 1.9 1.9 1.4 0.5 0.9 0. 5 0. 9 0.5  14  2.6 2.6 13.2 13.2 2.6 2.6  5.3  12.3  36. 8  5. 3  5.3 2.6  10.8  3  7. 9  19  50.0  Unknown Causes  126  59.4  Grand Total  212  100.1  38  100.0  51  not  associated  with  obvious  predators,  w h i s t l e s c o u l d have some other f u n c t i o n alarm  calls.  Bopp  by  emphasized marmots  in  Armitage  addition  However t h i s  the  cause f o r them.  (1962)  and Barash  The  individuals  has  been  reaction  of  same whether or not I was able t o a s s o c i a t e a T h e r e f o r e , I t h i n k t h a t the "unknown" causes o f p r e d a t o r s that I  d i d not see or common d i s t u r b a n c e s such as s l i d i n g hail,  being  (1973b), both of whom  many of the w h i s t l e s I heard were due t o e i t h e r  wind,  to  function  the warning f u n c t i o n o f t h i s c a l l .  was  i s possible that  (1955) i n t e r p r e t e d the w h i s t l e s of Harmota  marmota as t e r r i t o r i a l c a l l s . disputed  it  or  moving  fog  patches  that  snow,  rain,  alarmed  certain  (Table I V ) .  I r a r e l y knew which marmot w h i s t l e d but i n each case when I d i d the i n d i v i d u a l was  already  at  a  burrow  entrance.  Upon  h e a r i n g a w h i s t l e marmots u s u a l l y ran to a burrow entrance or t o the  top of a rock t h a t had a burrow  ran and entered a burrow emerged  from  the  burrow  burrow entrance marmots disturbance.  directly;  Marmots  in  fact,  not  about  for  emerge is  until  from  unknown.  i n v o l v e s some r i s k entrance  frequently  the  Once at a  cause  of  adaptive  to  the  burrows  keep  any  i n s i g h t and a v o i d , i f p o s s i b l e , the r i s k of having t o  subsequently situation  they  immediately enter t h e i r  when d i s t u r b e d presumably because i t i s predator  Marmots r a r e l y  at the sound of a w h i s t l e .  looked do  underneath.  the  because  a  burrow  Emerging  when from  predators  may  the a  above  burrow wait  at  ground  definitely a  burrow  occupant emerges. , Coyotes ( C a n i s 1 a t r a n g )  have been observed c a t c h i n g Spermophilus columbianus  (Don  Bowen  52  personal  communication) and M • c a licj at a ( I . McT.  communication), catching  S.,  and foxes {Vuljjes undulatus  vuljaes)  {Gordon  genus Pedetes, a r e presumably  tactics. emerging  They from  apparently their  the  O c c a s i o n a l l y some at  confound  and  by Waring  (1966), Barash  in  only  predators {Vaughan a  few  by  1972). minutes  apparent.  marmots  what  that  did  The  not  appear  intensity  to  react  to  from where they  {loudness)  of  the  not the w h i s t l e d u r a t i o n or i n t e r v a l as suggested  action  (1973b) , and Gray  (1975),  marmots would take.  not measured i n the f i e l d ,  whistles  awaiting  a l l , o r they merely looked around  whistle,  was  rodents  s u b j e c t e d to the same hunting  at t h e i r burrows  happened t o be at the time.  determine  observed  p r e d a t o r s had disappeared and w h i s t l i n g had stopped.  No " a l l c l e a r " c a l l was  whistles  been  African  burrows with a great l e a p  Marmots u s u a l l y remained after  have  Haber p e r s o n a l communication)  u s i n g t h i s technigue.. S p r i n g hares, south the  Cowan p e r s o n a l  appeared  Although  intensity  i t was obvious from l i s t e n i n g  intensity varied greatly.  to  to  More i n t e n s e w h i s t l e s  r e s u l t e d i n more marmots running to burrows.  53  Long w h i s t l e s were more o f t e n a s s o c i a t e d  with  terrestrial  d i s t u r b a n c e s , and short w h i s t l e s were more o f t e n a s s o c i a t e d aerial  with  disturbances. number of w h i s t l i n g seguences d i s t u r b a n c e type aerial terrestrial xz=16.81  Two  whistling  there were However,  of a medium  long  and  25 3 n=4 7  short  whistles  in  the  c a l l o f each sequence  was  considered.  Four  were dropped from t h e a n a l y s i s because the c a l l s  were  length.  Island  detailed  observations  marmots r e a c t e d t o alarm  calls.  not d e t e c t any d i f f e r e n c e s between responses t o long whistles.  sequence.  i n s t a n c e s were c o n s i s t e n t with the above r e s u l t s  I d i d not r e c o r d very Vancouver  5 14  seguences were dropped from the a n a l y s i s because  i f o n l y the f i r s t seguences  short  p<0.001  df=1  both both  long  of  the  way  Thus I could and  short  54  Keeaws:  Vancouver  Island  marmots  p r e v i o u s l y been recorded was The  a  faint  ±39  two-syllable  Hz  to  harmonics, both The  mean  a  call  vocalization  of a keeaw changed i n two stages  less intense of  This  which sounded l i k e a "kee-aw".  1109 ±57 Hz ( F i g 14).  duration  sound t h a t has not  f o r other marmots.  fundamental frequency  1912  produced  than  the  from  There are at l e a s t two  fundamental  frequency.  a keeaw c a l l was 0.29 ±0.016 s.  Keeaws  were u s u a l l y given i n a long s e r i e s t h a t v a r i e d c o n s i d e r a b l y length.  On two o c c a s i o n s only s i n g l e keeaws were given but the  mean number o f keeaws per sequence was 102 ±27. about 900 keeaws were given i n 60 The  interval  calling  in  between  keeaws  minutes  averaged  seguences the i n t e r v a l between  by 3.8  calls  On one o c c a s i o n one  individual.  ±0.15 s.  In long  increased  toward  the end of the sequence.  Keeaws  were  frequently  associated  t h e r e f o r e a l s o with w h i s t l e s (Table IV and keeaws  were  predators represent  Upon  usually  had  left  the  area.  keeaw  calls  Keeaws  there.  15).  Marmots  therefore  or low i n t e n s i t y  However, and t h e  seemed  alarm.  to  gradually  a  rock  resumed  or  burrow  their  keeaws continued.  and  previous  a c t i v i t i e s w i t h i n a few minutes a f t e r the c a l l i n g s t a r t e d , when  to  some marmots d i d not appear t o  r e a c t a t a l l , but many marmots ran rested  Fig  given a f t e r w h i s t l i n g had stopped  a s t a t e o f "uneasiness"  hearing  with d i s t u r b a n c e s and  even  In a l l but one i n s t a n c e only one animal  55  F i g u r e 14. R e p r e s e n t a t i v e sonograms o f 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. 15001000. 5000* 0*0  0.1  0«3  TIME  0.5  (SEDLTCE)  ~t  0.7  r  0>9  1  56  F i g u r e 15.  Frequency with which w h i s t l e s and keeaws together and s e p a r a t e l y with and without d i s t u r b a n c e as a s t i m u l u s  occurred a known  L i g h t numbers i n d i c a t e the number of o b s e r v a t i o n s dark numbers i n d i c a t e the percent of the t o t a l v o c a l i z a t i o n sequences  56a  57  gave keeaws at any one  Rapid  time.  Chirps  On  one o c c a s i o n when I approached a marmot colony  r a p i d s e r i e s of e g u a l l y spaced, very s h o r t w h i s t l e s  I heard  or  a  chirps.  I d i d not record t h i s c a l l , nor d i d I ever hear i t again.  Hisses  Individuals  in  live  approached the t r a p . with  its  mouth  traps  occasionally  When a h i s s was  open,  attempt to bite.„ The  crouched,  h i s s spans a  between 70 and 3000 Hz  " h i s s e d " when I  given the marmot faced  and wide  sometimes range  lunged  of  me  i n an  freguencies  ( F i g 1 6 ) . H a r m o n i c s t r u c t u r e , i f any, i s }  very weak.  Tooth C h a t t e r s  A Balph and  tooth 1966;  Banks  c h a t t e r denotes t h r e a t i n many rodents  Waring 1966,1970; Ewer 1968; 1973).  I  heard  c h a t t e r on only one o c c a s i o n .  a  Barash  Vancouver  T h i s occurred  the animal, an a d u l t male, f o r t a g g i n g .  {Balph and  1973b;  Brooks  I s l a n d marmot tooth when I was  handling  I d i d not hear e i t h e r a  h i s s or a tooth c h a t t e r from f r e e ranging marmots.  58  F i g u r e 1,6. R e p r e s e n t a t i v e sonogram o f a h i s s Recording speed: 19.05 cm per s Playback speed: 19.05 cm per s Harrow bandwidth f i l t e r  FREOJENCY  (HZ)  59  Screams and Growls  "Screams" and "growls"  were o c c a s i o n a l l y heard  f i g h t s and chases but were not recorded, freguency  sound probably  with  a  during play-  A growl i s a b r i e f low  wide  range  Screams sounded l i k e long high freguency  of  frequencies.  growls.  SOCIAL BEHAVIOUR  S o c i a l Behaviour  I  Patterns  recognized  I s l a n d marmots. considered to  social  behaviour  patterns i n Vancouver  G r e e t i n g , a n a l s n i f f i n g , and p l a y - f i g h t i n g were be  behaved  in  Chasing,  mounting,  suckling,  a  13  reciprocal  acts  because  s i m i l a r manner when performing  social  alert,  avoidance,  grooming,  tail  play-fight  The both  interactants  these  behaviours.  raising,  invitation,  chasing were c o n s i d e r e d t o be n o n - r e c i p r o c a l behaviour  both  acts  lunging, and p l a y -  because  the  only d e s c r i b e s the a c t i o n of one of the i n t e r a c t a n t s .  d e s c r i p t i o n of scent marking i s  included  f o r convienience  and because of i t s s o c i a l  G££§£i22  Greeting  ( )• G  consists  in  this  section  significance.  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 ( r a r e l y ) s i d e of pattern fiariota  has  been  another  individual.  This  behaviour  d e s c r i b e d f o r a t l e a s t f i v e other s p e c i e s o f  (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 t h e most common s o c i a l t h a t I observed  AfiSi  behaviour  pattern  (Table V I ) .  sniffing  (AS). Anal s n i f f i n g c o n s i s t s o f two animals  s t a n d i n g together with t h e i r bodies p a r a l l e l while n u z z l i n g  the  anal region o f the other marmot.  Mounting other  from  (Mo). behind  Mounting i n v o l v e d one animal s t r a d d l i n g the with  i t s f o r e l e g s and p l a c i n g i t s v e n t r a l  s u r f a c e i n c o n t a c t with the d o r s a l s u r f a c e of another. observed other  the  marmot  Suckling  never  d o r s a l animal t h r u s t i n g or b i t i n g the back o f the as  M. f l a v i v e n t r i s  I  was  seen  in  the  sexual  behaviour  of  (Armitage 1965) and M. olyjngus (Barash 1973b).  (S) and S o c i a l Grooming  were observed o c c u r r i n g o n l y between  (SG). Both of these a c t s adult  females  and  their  infants.  .Avoidance was  sure  (Av). I recorded avoidance behaviour o n l y when I  that an animal's departure was i n response t o another  individual.  T h i s occurred when a marmot e i t h e r ran away  during  an i n t e r a c t i o n o r moved away from an approaching marmot once the latter  was w i t h i n 3 m.  Avoidance does not i n c l u d e animals that  were f l e e i n g d u r i n g a chase.  Alert appeared  (Al).  A l e r t behaviour was performed  by marmots  that  t o be uneasy about the approach o f another i n d i v i d u a l .  The a l e r t e d animal watched the approaching marmot from  a  rigid  61  crouched  stance.  £1131113  (*•) •  Lunging c o n s i s t s of t h r u s t i n g the  foreward toward another marmot, o c c a s i o n a l l y  l a i i Raising consists the  tail  until  of up,  the  (TR).  making  f r o n t paws contact.  T a i l r a i s i n g i s a graded d i s p l a y  which  a marmot e r e c t i n g t h e h a i r on the t a i l and noticeably  arched.  fluffed t a i l  The  tail  may  raising  be r a i s e d  further  l i e s f l a t along the back o f the  animal.  When t h i s d i s p l a y i s given the t a i l i s u s u a l l y moved very or held  motionless at any  point  within  t h i s range of  All  t a i l r a i s i n g d i s p l a y s were performed at the  of  four  behaviour  patterns:  a l e r t , or p l a y - f i g h t i n g .  either  However, not  consider  positions.  same time as  greeting,  anal  sniffing,  raising.  Thus,  Very s i m i l a r t a i l r a i s i n g behaviour was (1973b:184,198,  (1967:44 and Fig  5) and  is  very  50)  and  Fig  23)  i n M. c a l i g a t a , and  Waring  (1966: 181)  d i f f e r e n t from the  i n M.  by  in  from the  arched  tail  1974:243).  "tail  M.  oly.S£H§»  flaviventris.  i s r a i s e d up and  H.  by  by  Gray  (1962:325  and  This  marmota  that  (Koeing  1957)  which  waved from s i d e to s i d e involves  behaviour  swirling  flaviyentris in  T a i l r a i s i n g i n H. Sonax a l s o  pattern.  observed  the r a p i d pumping and  f l a g g i n g " of  movement (Bronson 1964:471).  also  Armitage  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. and  one  t a i l r a i s i n g to be an o p t i o n a l component of each  of these behaviours r a t h e r than as a separate behaviour  Barash  one  a l l of the occurrences of  these behaviours were accompanied by t a i l could  slowly  the  (Armitage  rapid  tail  62  Chasing  (C)  and  Plav.-cjiasing  a g o n i s t i c encounter t h a t  differed  (PC).  from  Chasing  what  I  was  termed  an  play-  c h a s i n g i n the f o l l o w i n g ways: 1.  P l a y chases were slower and s h o r t e r , u s u a l l y l e s s than  7-10  metres. 2.  Play-chases were always s i l e n t whereas a g o n i s t i c chases were  o c c a s i o n a l l y accompanied by w h i s t l e s , s g u e a l s , or growls. 3.  Play-chases always ended with the  feeding  close  together  interactants  were  or  or i n t e r a c t i n g i n a n o n - a g o n i s t i c way.  A g o n i s t i c chases were r a r e l y f o l l o w e d by other interactants  resting  usually  separate  acts  since  a f t e r the chase  the  (Fig 17).  Chasing and  p l a y - c h a s i n g always caused  thus i t was  not necessary t o d i s t i n g u i s h " f l e e i n g " as a separate  behaviour  pattern.  Play-fighting rising  the other marmot to f l e e ,  up  on  (PF).  P l a y - f i g h t i n g c o n s i s t s of two  t h e i r hind l e g s and pushing with t h e i r f o r e l i m b s  a g a i n s t the other marmot's c h e s t or s h o u l d e r s . has  also  been  H. f l a y i v e n t r i s  described  that  1974,  (PI) .  appeared  performing t h i s behaviour posture  M.  This  olympus Barash  behaviour  (Barash  1973b),  1973a), M.  caljgata  marmota (Barash 1976b).  Piai-figjlt Invitation signal  for  (Armitage 1973,  (Barash 1974b), and M.  visual  marmots  was  A play-fight invitation to  indicate  prepared  to  that  the  play-fight.  was  a  marmot This  ranged from simply r a i s i n g the s h o u l d e r s t o r a i s i n g the  whole body i n t o a v e r t i c a l p o s i t i o n on the back l e g s .  63  Scent made  Harking  (SH) . , V a n c o u v e r  l o n g sweeping  motions  were a t t h e e n t r a n c e s t o deposits  scent  with t h e i r  burrows.  i n t h e form  All  marking  North  was  American  have f a c e g l a n d s s c e n t , marking  usually and  performed  and  Gray  U.  olvmp.us, B a r a s h  Dominance  M.  a s an  H.  presumably face  individual  indicated  dominant  marmot i n a c h a s e  Lunging  was  behaviour.  use  1967:48; M. cited  them  for  marmota,  Koeing  i n R a u s c h and  Rausch  and  Rausch  flavj,ventris,  also  the  dominant was  marmot o f an  the  animal  1971:90-91;  Armitage  1976b).  characteristic  o f dominant  characterized  interaction. avoidance  in  c o n s t r u c t e d as d e s c r i b e d Table  Va.  >  two-year-old  by  i n the form,  females  >  The  chasing.  animals since i t and  avoidance  individuals  of  an  on c h a s i n g , l u n g i n g , and  I t i s apparent  dominance h i e r a r c h y e x i s t e d  the  Flight  subordinate  A dominance m a t r i x b a s e d  was  presented  the  lunging,  interaction.  doing  c a u s e d o t h e r marmots t o move away.  therefore,  females.  but  Relationships  clearly  females  glands.  interaction,  T h r e e b e h a v i o u r p a t t e r n s , c h a s i n g , a v o i d a n c e , and  usually  that  E u r a s i a n marmot s p e c i e s  broweriRausch  1973b: 184;  rocks  behaviour  social  1967  frequently  against  R a u s c h 1971:90) and  (M. c a l i g a t a .  monax and  This  after  1957:519; A s i a n marmots, B i b i k o v 1971:90; M.  cheeks  some, i f n o t a l l ,  (Rausch  marmots  o f s e c r e t i o n s of t h e i r  Marmots s o m e t i m e s s c e n t marked scent  Island  Brown (1975) from adult  yearling  and  these data that males males >  >  is a  adult  yearling  64  Table  V.  Dominance matrices o f Vancouver I s l a n d marmots  (a) . Dominance Matrix Occurrence of Chases, avoidance, Classes  Dominant  AM AF ""'"\;.'?2F' ' V ''•'.-."''2?'; ' . . YM :  Subordinate AF 2F 2?  AM*  3 2  1  sex  YM  2  1  YF  Row Column Totals Totals  Y?  2 10 1 2  2 2  ::  1  '' '''T''~ ' "Yt i!  :  :  c l a s s e s of  Based on the Freguency o f and Lunges Between Age:sex  :  i  l  7  age and  1  2  Y?  7 24 4 0  1 4 7 2 1  0  14  0  6  (b) . Dominance Matrix Based on the Freguency o f Occurrence of Eight Behaviour P a t t e r n s (C, Av, A l , L, PI, TR, M, SG) Between Age:sex C l a s s e s  Dominant  AM AF A? 2F 2? YM YF Y? I?  Subordinate A? 2F 2?  AM  AF  2  23 2  1 1  3  1  3 1  1  * KEY AS IH TABLE I I I  8 13  4 2  YM  YF  Y?  2  7 10  3 3  2  2  3  3  I?  3 3  Row Column Totals Totals  48 36 3 14 4 0 0 0  1  6 29 3 21 6 4 22 9 6  65  When respect was  the  to  other  the  that  social  indicating  marmots  performed  dominance  matrix  Table  six)  Vb.  of  most  in  likely  Ho  between  to  the  or  raising  these  exceptions  age  hierarchy,  and  where  sex  it  invitations, also  status  play-fight  and  subordinate  behaviour.  behaviours  {reversals),  between  Table Va,  performed  mounting,  with  is  presented  five  {possibly  classes  variation  The  that  were  would  be  occur.  relationships  number  greetings anal  terminate  any  (X =0.60,  n~15)  2  11  of  in  were  marmots  tail  8  analyzed  play-fight  alert,  and  and  on a l l  dominance  patterns.,  initiated n=17),  alert  occurred  dominance  behaviour  grooming,  Of the  the  and  were  indicated  raising,  Dominant  based  these  adjacent  tail  grooming,  social  patterns  relationships  behaviours.  invitations,  in  behaviour  dominance  apparent  mounting,  ten  There of  more than  were  dominant  {X =0.78,  apparent  no  significant  play-fights  {X =2.00, n = 1 8 ) . 2  greetings  (X =1.00,  d i d dominant  among  and s u b o r d i n a t e  n=33) ,  2  sniffs  were  2  marmots.  the  other  differences marmots  or  that  (X =0.06, 2  Subordinates n=25)  5  did  not  play-fights  66  The  Frequency  of S o c i a l  I observed social  a total  interactions  Vancouver I s l a n d observed  o f 785  over  marmots.  between  the r e l a t i v e  Behaviour  age-sex  not  two  The  freguency  interactions,  the  just  observed  in a specific  Dominance  Relationships).  all  most s t r i k i n g  age-sex  each  or  Island  used  patterns  that  I  characteristic  behaviour  the  same  behaviour  varied  among  relatively  rare.  degree  not  be  independent  of these  can  adult  groups.  the  usually  dyad  behaviour  of  whole  classes  on  that  patterns  in  play-fighting by  Vancouver  a l l  behaviour  study  the next  freguency  and  were  most  that  commonly  o f t h e o t h e r 10  acts  but  a l l  they  were  among a g e - s e x  statistically  greetings  was  i n Table VII i s  65%  VI  non-reciprocal  p a t t e r n (see s e c t i o n  of s i m i l a r i t y  compared  i.e.  females  was  in  p a t t e r n s used  for  and  within  o f a l l s i x age-sex c l a s s e s  age-sex  The  are  and  over  The  different  VII)  males  raising  pattern.  (Table not  accounted  observed  Tail  was  data i n Table  G r e e t i n g and  behaviour  of the behaviour  I recognized. used  They  act  f r e g u e n t l y each  f e a t u r e of the d a t a  most common s o c i a l marmots. ...  587  observed  occurred  The  recipient  a p p r o x i m a t e l y t h e same p r o p o r t i o n s . were t h e  I  each  behaviour  shows how  social  classes  that  that  i n Table VII.  actor  the t a b l e  summers  in  c l a s s e s i s p r e s e n t e d i n T a b l e VI  freguencies that  indicate  The  behavioural acts occurring  the  age-sex c l a s s e s i s p r e s e n t e d do  Patterns  because  occurring  increase the greeting  classes the  between  freguency of  data adult both  <0  Table  V I . The f r e q u e n c y  DYAE  *(!:AF AM:2F AB: YF AR: YM AM:? AF: AF A F: 2 F AF: YF AF: YM AF:I AF:? 2 F: YF 2 F : YM 2F:? Y F : YF Y F : YM YI: ? YM:? 1:1 I:? ?: ? TOTAL  G  33  7 6 4 21 41 7 8 11 18 6 3 8  1 1 4 1 16 2a . 70 290  AS  c f o c c u r r e n c e o f each  C  AV  4 1 2  2  2  5  6 2  2  6 1  a 1  9  1  1  18  2  2 9  4 1  1  6  11  a  2  1 2  1  3  6  28  27  60  28  5  ii  behaviour  PC  PI  1  10  M = Bale 2 = T w c - y e a r -o l d Y = Yearlinq f = Female ? = Sex and I = Infant age unknown G = Greeting' S = Suckling A = Adult PI = P l a y - f i g h t Invitation  3  1  1  1  1  3  31  13 4 118  2 2 30  88  217  36  C Av PF TR Al  = = = = =  Mo  SG  S  2 1 1 3  n u n  1 1  L  between age-sex  3  5  7 1  pattern  1 1  4  1 1 6  1  PF  5  1  1  TR  16 8 9 5 7  2 3  Al  social  Chasing Avoidance P.lay- f i q h t i n q Tail Raising Alert  2  2  3  5  TOTAL  78 26 24 11 57 2 69 24 13 19 52  10 9 15 6  1  7  classes  3  7 1 32 1  2  7  7  PC L Bo SC AS  = = = = =  35  292 3  5  Play-chasing Lunging Mounting S o c i a l Groominq Anal S n i f f i n q  785  OO  Table  VII. Percentages  of social  behaviour  patterns  p e r ane-sex  class  ID  AGI-SEX ciass  G  AS  c  AM  36  6  5  AF  46  4 .  2F  50  YF  33  YM  46  I  57  3  •»  29  3  10  3  AVERAGE  37  3  8  4  Av  Al  TR  PF  4  3  19  23  10  4  2  12  1 5  2  2  6  4  10  22  6  11  4  9  30  3  5  5  35  3  25  5  1  11  34  9  <1  1  1  1  1 1  28  5  1  1  1  3  1  PC  <1  PI  L  Mo  2  1  2  1  2  2  3 1  3  SG  1  S  2  1 4  3 3  <1  4  1  -  M = F = ? =  Male 2 = Two - y e a r - o l d Female Y = Yearling Sex and I = Infant a g e unknown G = Greeting A = Adult S = Suckling EI = P l a y - f i g h t I n v i t a t i o n  C Av PF TR Al  = = = = =  Chasing Avoidance Flay- fighting Tail Raising Alert  PC L Mo SG AS  = = = = =  Play-chasing Lu n g i n g Mounting S o c i a l Grooming Anal Sniffing  BOW TOTAL  TOTAL ACTS  101  196  101  259  100  129  101  80  100  37  101  118  102  751  101  1570  69  The s i m i l a r i t y i n the behaviour of age-sex c l a s s e s whole  does not r e v e a l anything about  o c c u r r i n g between compare  specific  dyads.  It  p a t t e r n s occurred between dyads because amonq  Behaviour). of  individuals  (see  was  not  the  possible  section  on  of  any  given  to  behaviour  observation  times  Hates of S o c i a l  However, what can be compared are  occurrence  the  the nature of i n t e r a c t i o n s  the a b s o l u t e f r e q u e n c i e s with which d i f f e r e n t  differed  on  the  frequencies  act as a p r o p o r t i o n of the t o t a l  number of a c t s .  In order to t e s t the  relative  different  frequency  dyads  independence.  f o r s i m i l a r i t y between  (Table  with VI)  which  using  acts  a  1)  chasing,  between  test  for  less  than  one,  I  Armitage  1962,  1976a), and 2) g r e e t i n g and a n a l s n i f f i n g , because  of the  than  twenty  I  used  T h i r t e e n of 15 comparisons thus  occurred  a l e r t , avoidance, and l u n g i n g , because o f acts  cohesive nature of these a c t s . less  compared  When i t was necessary t o lump t h e f r e q u e n c i e s of  the a g o n i s t i c nature of these 1973,  I  Chi-sguare  c e r t a i n behaviours t o avoid expected values lumped:  dyads  (see  also  I f the t o t a l number of a c t s  the  F i s h e r Exact P r o b a b i l i t y  were not  significant  (Table  was test.  VIII),  i n d i c a t i n g that the behaviour p a t t e r n s were independent  the i n t e r a c t a n t s . infant : infant  That  i s , the  interactions  behaviour were  behaviours used between i n f a n t s and behaviour  patterns  used  in  i n t e r a c t i o n s d i f f e r e n t from those  not  patterns different  non-infants adult used  nor  used  adult  in  from  the  were  the  male : y e a r l i n g in  of  male  male : a d u l t  70  Table  VIII.  Comparisons o f t h e r e l a t i v e f r e g u e n c y w i t h which social 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 between d i f f e r e n t age-sex c l a s s e s  COMPREISON  AM: AF AM: AF AM: YF AM: Y F AM: YF AM: YM AM: 2F AF: 2F AF: YF AF:2F AF: YF 2F:YF 2F: YM YF: YF  1:1  * = = M = F = A = 1  AM: 2F  - AM: YF - AM: YM - AM: 2F - AF: YF -• AF: YM - AF: 2F - AF: YF - AF: YM - AF: YM  -  -  -  YF: YF Y F : YF 2F: YF YF: YM I:NI  P<0.05 F i s h e r Exact Male Female Adult  n  X2  df  P  104 10 2 35 50 48 24 95 93 37 82 30 16* 19» 9* 67  2.76 3.75 .49 .35 11.55 1.62 5.49 25.96 4.6 8 .17 4.80  4 4 2 4 4 2 3 3 2 3 2  2.59  2  .60 .44 .78 .98 .02* .45 .14 <.01* .09 .91 .09 .96 .42 .21 .27  P r o b a b i l i t y T e s t , N<20 2 = Two-year-old I = Infant NI = N o n - i n f a n t  71  female or a d u l t male : y e a r l i n g female i n t e r a c t i o n s . for  the  comparisons between a d u l t females and  and other dyads were s i g n i f i c a n t VIII,  AF:2F and AH: YF)  VIII).  There was  yearling  more  females  than  at the 5%  or the 10% l e v e l aggression there  example, g r e e t i n g s made up  a  was  (YF: YF and adult  between  much  yearling  level  between  higher  Chi-sguare  other  {*  females  in  Table  AF: YM,  Table  females  and  dyads.  For  proportion  of  the  i n t e r a c t i o n s between a d u l t females and two-year-old females than between the  adult  females and y e a r l i n g females  o p p o s i t e t r e n d was  e v i d e n t with chases  A few dyads were s u f f i c i e n t l y pattern  (59% vs 29%), while  (4% vs 25%).  different  from  general  of behaviour t h a t I observed between age-sex c l a s s e s as  a whole, t h a t s t a t i s t i c a l t e s t s were not r e g u i r e d . were  the  never  seen t o i n t e r a c t  Adult  males  with other a d u l t males even  though  t h e r e were two i n d i v i d u a l s present on the Haley Lake study throughout  the whole of 1974.  Adult females i n t e r a c t e d on only  two o c c a s i o n s , both o f which were chases. significantly non-infants  less  a g o n i s t i c behaviour  (X2=13.59,  area  p<0.0l, df=1) .  Infants  experienced  (C, A l , Av, L) than d i d  72  I n t e r a c t i o n 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 this  study  consisted  o f only one behaviour p a t t e r n .  during However,  21%  (124 o f 587) o f a l l i n t e r a c t i o n s c o n s i s t e d of a sequence  two  o r more a c t s .  was  2,6  acts  of  The average l e n g t h o f an i n t e r a c t i o n sequence  (SE=0.J2, range= 2-7). There were no s i q n i f i c a n t  d i f f e r e n c e s amonq age-sex  classes  or  specific  dyads  i n the  average number of a c t s per i n t e r a c t i o n .  Greeting  was  usually  i n t e r a c t i o n sequence. seguences,  The occurred  relative  30%,  behaviour  of  and  of  an  a l l interaction  seven  other  acts  17% of the seguences.  freguencies  with  which  two-act  seguences  17. , Since t a i l r a i s i n g always  at the same time as some other a c t , r a t h e r than  or a f t e r i t , i t was omitted 17.  began  i s i l l u s t r a t e d i n Figure  occurred  initial  G r e e t i n g s began 53%  play-fighting  i n i t i a t e d the remaining  the  before  from c a l c u l a t i o n s r e l a t i n g to F i g u r e  Host i n t e r a c t i o n s proceeded from  a  greeting  to  a  play-  fight.  Some  behaviour  p a t t e r n s occurred  much more o f t e n than they of  27  anal  sniffs,  in interaction  occurred as s i n g l e a c t s . ,  26 o f 36 play-chases,  f i g h t i n t e n t i o n s occurred  i n sequences.  seguences Twenty-five  and a l l seven play-  73  F i g u r e 17.  Temporal occurred  context  with  which  two-act  seguences  The width of the l i n e i s p r o p o r t i o n a l to the t h e freguency with which t h e behaviour sequence occurred The narrowest l i n e r e p r e s e n t s 2% o f a i l seguences 26% of a l l sequences proceeded from a g r e e t i n g to a play-fight  74  Rates  of  Social  Field  observations  investigator an  Behaviour  egual  to  observe  amount  performed  is  behaviour  of  is  differences  species,  whereas with  behaviour  The  in  each  among  of  the  fighting  which  behaviour  all  X.  behaviour (Fig  rates  can  the  group o f  animals  for  behaviour  be  used  to  is  describing  to  compare  the  individuals  compare  patterns  pattern  for  used  between  of  than  the  occurred  did  and  relative  within  class,  with  greeting and  of  the  class,  Vancouver  in  the  rates  IX)  than  adult rates.  and  and  colony  there  two  to  of  greeting  yearling  in  the  of  two  the  were  behaviour in  often  for final  patterns  freguencies high  females  had the  most  class  marmots.  p l a y - f i g h t i n g were  Yearlings  had  colonies  Island  was  between  age-sex  obtain  two-year-old  females  somewhat  corresponding  results  rates  (Table of  the  age-sex  variation  varied  A l l marmots  the  each  Rates  patterns  18).  averaged  classes  rates  and  ^The Table  only  I  VI).  highest  chases  can  more  age-sex  Rates  be  a  allow  parameter  counts  interaction  agezsex  (Table  useful  behaviour  a n d two  one.  was  more  each  that  the  of  behaviour  There  The r a t e  in  greeting  estimate  all  one  colony  animal or  rarely  individuals.  rates  colonies higher  of  a  behaviour  frequency.  absolute  freguency  each  time.  therefore  than  of  highest  in  having play-  involved  in  presented  in  lunges.  rates  of  Two v a l u e s  behaviour are  patterns  presented  per  where  dyad are  appropriate,  for  non-  75  F i g u r e 18.  A one  comparison and two  of i n t e r a c t i o n r a t e s between c o l o n i e s  06  2.0 r  H • I  c  colony one colony two  •1.5  in c  o y  1.0  u (TJ (D  £  0.5  A M AF 2F YM  Greeting  AM AF 2F YM  Play Fighting  AM AF 2F YM  Agonistic (c, A V , A I , u  AM AF 2F YM  Tail Raising  76  Table  IX. Interaction rate per thousand hours  Age:Sex :lass  age-sex  class cf  Behaviour Pattern Av Al TE  G*  AS  c  AH  488  513  0  99  160  AF  1225  32  56  32  2F  1005  23  22  IF  388  0  Yfl  606  0  * KEY AS IN TABLF VI  behavioural  acts per  PF  PI  L  Ho  30 1  138  74  0  24  0  80  265  8  32  74  45  203  44C  271  102  0  79  388  97  0  0  583  0  291  0  0  0  0  101  303  0  0  0  77 X. Interaction rate per dyad per behaviour per thousand hours  DYAD  G*  AS  C  Av  Al  TR  PF  AM: AF AM:2F AM: YF AB: YM AF: AM AF: AP AF: 2F A F : YF A F : YM 2 F : AM 2F.-AF 2 F : YF 2 F : YM Y F : YF Y F : YM YM: AF YM: YF  95 78 32 38  12 4 12  2  4  1 70  -  -  49 76 20 35 2  28 26 42 30  37  27 30 61  9 15  _  258 30 228  -  58 76 20 8  -  9  10 4 14 7 41  -8  12 6  3 11 15  -  * KFY AS IN TABLE VI  14 2  9 62 73 _  PI  L  7 38  78  'reci.pr.pcal  behaviour  patterns,  but  d i s c u s s i o n of Dominance R e l a t i o n s h i p s , r  reversals  i.e.  were highest highest  yearling  f o r g r e e t i n g and  greeting  females and highest  of  For  agonistic most dyads  there  chasing  and  behaviour (9 o f  was  The adult  (Fig  the  higher  i n June than i n a l l other  Greeting  r a t e was  increases  morning, and  of  just  few  emergence  marmots  The  dyad with to  the  yearling  highest  in  June  September (Fig  behaviour  patterns much  months.  r e l a t i v e l y constant after  Rates  two-year-old  a g o n i s t i c acts v a r i e d the most, being  j u s t before  ( F i g 21) .  a  only  19).  were  r a t e s v a r i e d l e s s than other  rate  the  12) the g r e e t i n g r a t e s exceeded  r a t e s of a l l behaviour p a t t e r n s  Greeting  in  a d u l t females.  y e a r l i n g males.  while  night  were  g e n e r a l l y d e c l i n e d through J u l y , August, and  20).  with  noted  between a d u l t and  the r a t e s of a l l a g o n i s t i c a c t s combined  The  was  p l a y - f i g h t i n g i n most dyads.  r a t e s occurred  a d u l t females and  rate  females.  males  as  entered  throughout  from the  the  day,  the burrow i n the burrow  for  the  79  F i g u r e 19.  G r e e t i n g and  agonistic  interaction rates  per  dyad  79a  INTERACTION RATE PER THDLJSANO ANlMAL-t-OJRS  6  H >  >  -n I  1  8 » 6 - S 8 S 8 8 § g S ' B S B S 1  1  1  h  H  1  1  1  1  1  1  1  1  80  F i g u r e 20.  Variation  in interaction  rates  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  F i g u r e 2 1 . V a r i a t i o n i n the g r e e t i n g r a t e throughout the  day  81a  GREETINGS PER ANIMAL-HDUR i  o  ru  o -P  o  o h  o P  H P  K  K p — P  ^  -  F £  r  u  r  P — p — P  u  r P  u f  82  D i s p e r s i o n , T e r r i t o r i a l i t y , and Scent  On The  colony  two,  Marking  a l l f o u r marmots had t h e same home range.  s i z e of t h i s area was approximately  the  dispersion  was  3 Ha.  o f marmots was more complex.  a l a r g e amount of home  range  overlap  On  colony  In May 1974 there  among  a l l animals.  Most of t h i s o v e r l a p occurred i n t h e area with steep c l i f f s 22). it  Until  one,  (Fig  mid-June t h i s was t h e o n l y area f r e e of snow, thus  was t h e only area where food was a v a i l a b l e .  By the middle o f June, most of the snow had melted from the colony and marmots f e d on p l a n t s t h a t were growing on the c l i f f s much l e s s f r e g u e n t l y . between  the home  decreased. they  At  ranges  this of  that  were  half  a d u l t male, #4, emigrated  e a r l y June and the remaining  adult  male,  overlap  separate  2.2 Ha of t h e colony.  the c o l o n y  and  Ha  ( F i g 22).  However,  until  a  t o colony one at t h e end of adult  female  #12  on the  S h o r t l y a f t e r the a r r i v a l o f a d u l t  drastically  reduced  h i s home  u n t i l i t d i d not o v e r l a p a t a l l with t h a t o f #17 (Fig 24). through  adult  adult male, #13, moved over the  #17, immigrated  male #17, a d u l t male #13  July  and  from colony one t o c o l o n y two  June and occupied a s i m i l a r area to upper  of  (Fig 23).  whole colony, an area of about 4.5 third,  completely  of  female #12 occupied the upper h a l f  in  amount  way f o r t h e d u r a t i o n of t h e summer., Adult  female #15 occupied the lower  One  the  t h e two a d u l t females g r a d u a l l y  By J u l y t h e i r ranges  remained  time  range From  the animals hibernated i n September there  83  F i g u r e 22. Home ranges o f t h e two adult females (#12 and #15) i n May 1974, and the two a d u l t 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 l o c a t i o n s of scent markings made by a d u l t male #13 before the a r r i v a l of a d u l t male #17 (n=3). This figure i l l u s t r a t e s the same area shown i n F i g 2B.  83a  84  F i g u r e 23. Home ranges o f four female marmots on the Haley study area i n J u l y 1974  Lake  Dots i n d i c a t e t h e l o c a t i o n s of a l l scent marks made by a d u l t female #12 (n=4). C i r c l e s indicate the locations of a l l s c e n t marks made by two-year-old female #9 (n=5). Triangles i n d i c a t e t h e l o c a t i o n s o f a l l s c e n t marks made by a d u l t female #15 (n=5) . Arrows i n d i c a t e t h e l o c a t i o n s of chases. #14 i s a y e a r l i n g female. #17 i s an a d u l t male.  85  Figure  24. Home ranges o f the two a d u l t males on the Haley study area i n August 1974 Dots i n d i c a t e t h e l o c a t i o n s of a l l scent marks by a d u l t male #13 i n June, a f t e r t h e a r r i v a l o f male #17 (n=2). T r i a n g l e s i n d i c a t e t h e l o c a t i o n s (n=13) o f a l l marks (n=17) made by a d u l t male #13 i n J u l y . Squares i n d i c a t e t h e l o c a t i o n of a l l scent marks by a d u l t male #13 i n August (n=1) . Circles i n d i c a t e the l o c a t i o n s (n=6) of a l l marks (n=12) made by a d u l t male #17.  Lake  made adult scent made scent  85a  86  were c l e a r l y two main areas on t h e colony. occupied  Yearling  female  part  both  of  two-year-olds,  areas  ( F i g 23).  analysed  causes  occupants  colony  a  #15  Brown  territory  markings. ,  determine  and  being  my  I s l a n d marmot females a r e  Adult females Both  of  these  #12 from the area normally area  Orians  a defended  (Fig 23).  Both  (1970)  accept  the  area and e x p l i c i t l y  1.. a c t u a l defense such as chasing  2. performing  Thus  the  s c e n t marked w i t h i n t h e i r areas o f  d e f i n e defense as being e i t h e r  scent  to  occasions.  chasing  occasionally  away an i n t r u d e r , or  one  t h i s p a t t e r n of d i s p e r s i o n .  e x c l u s i v e use ( F i g 23). of  adult  used  seen.  used only by the occupants of t h e lower  concept  yearlings.  the b e h a v i o u r a l i n t e r a c t i o n s between  involved  females  three  occupied  of  One  #12 and #15 i n t e r a c t e d o n l y on two interactions  and  animal  of the upper and lower areas o f  adult  was  #14 was the only marmot t h a t c o n s i s t e n t l y  p e r i p h e r a l areas and was r a r e l y  proximal  area  by marmots #12 and #17 and t h e lower area was occupied  by animals #13, #15, two  I  The upper  data  identifying suggest  territorial  with  acts  such  as  t h a t a d u l t Vancouver respect  to  other  a d u l t females, but t h a t t h e i r t e r r i t o r i e s are s m a l l e r than  their  home range.  I  d i d not see any i n t e r a c t i o n s a t a l l between a d u l t  males  #17 and #13 i n s p i t e of the dramatic change i n the home range of #13 t h a t appeared There  could  have  to  be  been  directly  related  to  #17*s  some r a r e and s i g n i f i c a n t  arrival.  interactions  between these two marmots that I missed s e e i n g but u s u a l l y  these  87  two  animals j u s t avoided  enhanced  by  the  one another.  Avoidance  d e p o s i t i o n o f scent marks.  marked much more than a d u l t females. was  the  only  other  The  was  probably  Adult males scent  two-year-old  age-sex c l a s s of marmot observed  female to scent  mark. age-sex c l a s s  freguency  of scent marking  a d u l t males : a d u l t females two-year-old female  The  d i s t r i b u t i o n of #13*s  42 9 5  scent  marks  before  and  after  a r r i v a l o f #17  i s shown i n F i g u r e s 22 and  24.  marking  I observed  j u s t a f t e r the  of #17 his  that  ( F i g 25).  rate  was  done by #13  Adult male #4,  Host of the scent  on colony two,  of scent marking i n J u l y  a l s o have been due t o the presence  ( F i g 25).  also  increased  of the scent of another  j u s t immigrated to t h i s l o c a l i t y .  the ranges of #13  and  still  arrival  This increase  male s i n c e #4 had  #17  overlapped  may  adult  Even though  somewhat i n J u l y  each avoided the area enclosed by each others scent marks. area enclosed by scent marks was  the  they The  a l s o t h e area of maximum use of  each marmot.  , I analyzed the amount o f time that marmots spent not  feeding.  Adult  males moved s i g n i f i c a n t l y  age-sex c l a s s e s and a d u l t male #13 a d u l t males #4 and #17. spent range.  moving by a d u l t male #13 Patrolling  more than  moved s i g n i f i c a n t l y  The r e l a t i v e l y  large  moving but  amount  other  more than of  time  occurred as he p a t r o l l e d h i s home  i n v o l v e d moving around the perimeter of the  88  F i g u r e 25. Rates of scent marking by a d u l t #17 Numbers above bars i n d i c a t e marking bouts observed X i n d i c a t e s no data  males  the  #4,  #13, and  number  o f scent  Rate of Scent Marking (number/hr) o  O 0)  do  -T"  IV)  ro  o X  0)  > c C  CD CO  89  home  range  marking.  greeting Adult  other  male #4  marmots  d i d not  #13  I  think  t h a t a d u l t male #17  to  see  most  more  conspicuous  with  other  than  These o b s e r v a t i o n s marmot  males  males.  They occupied  feeding  are  female  #15  and  female #14  he  arrival  similar  to  and  2  adult  Vancouver  d i d not  against  first  of  those  adult  nor  arrived.  male  occurring and  #17  t r y to defend  #13,  #12  6 chases  interactions.  In  between #14  #13,  and  Some aggression  adult  (X  the  lower  or #14  and  adult These  between  adult  2  = 2.28,  the  upper  p>.25). of  and  the  d i d she r e a c t a g g r e s s i v e l y  to  As I mentioned e a r l i e r , y e a r l i n g lower  areas.  t r e a t e d q u i t e a g g r e s s i v e l y by  ( F i g 23)  be  #17.  c o n s i s t e n t l y moved between the upper and  #17,  Island  marks.  between #12  However, t h i s i n d i v i d u a l was and  X =18.3, p<0.01).  between adult female  the  #13,  Haley Lake colony  themselves  f i x e d areas of e x c l u s i v e use t h a t c o u l d  Thus a d u l t female #12  when  that  made  rocks  a l s o t e r r i t o r i a l with r e s p e c t t o other  before  i n t e r a c t i o n s were  allowed  by preceeding more of  (67% vs 28%,  to be defended by scent  #13  marmots  marmots  indicate  I n t e r a c t i o n s occurred male  adult  p a t r o l l e d l e s s often than  Moving  t h e i r movements with t a i l f l i c k s  considered  scent  of h i s home range from the tops of the  t h a t he f r e g u e n t l y r e s t e d on.  #12  contact  because the p h y s i c a l c h a r a c t e r i s t i c s o f the h a b i t a t  him  #1.7.  occasionally  p a t r o l h i s home range as o f t e n ,  probably because he never came i n t o males.  and  and  3 lunges out  area #15,  none of the were  both  of a t o t a l of  20  12 i n t e r a c t i o n s  chases  or  lunges.  d i d occur among marmots i n the lower area but i t  90  did.  not  appear  to  be  related  to  appeared as i f a l l occupants c o u l d move within t h i s  area.  s p e c i f i c l o c a t i o n s and i t about  freely  anywhere  91  DISCUSSION  VOCALIZATIONS  I n t e r s p e c i f i c Comparisons  The  vocalizations  s i m i l a r t o those However,  of  there  of Marmot V o c a l i z a t i o n s  of  Vancouver  M. c a l i g a t a  Island  and  marmots are very  H. olympus  are some important d i f f e r e n c e s .  (Table Barash  (1973b)  never heard w h i s t l e s from M. olyjijgus t h a t were as l o n g as commonly  given  given by  M. olympus  H«  :  by  M.  vancouverensis.  (Barash  1973b)  XI).  those  Rapid c h i r p s were often but  I  heard  them  from  vancouverensis only once.  The  keeaw  of M. vancouverensis corresponds i n M.  to a "queeuck" c a l l described  by  (Gray 1967)  Taulman  and  (1975).  the  The  "low  caligata  freguency  medium c a l l of M.  call" olympus  (Barash 1973b) although d i f f e r e n t i n sound s t r u c t u r e appears be  the  homologous  marmot c a l l i n g sounded  call  sequence  exactly  in that  l i k e a keeaw.  this species. I  heard,  In the one  the  first  d u r a t i o n and similar  to  interval that  of  of  medium  keeaws  M. oly_mj>us  uneasiness.  and  a l l three  few  (Table  XI).  calls  (1973b).  calls  is  calls toned The quite  Keeaw, queeuck, low  frequency and medium w h i s t l e c a l l s were a l l g i v e n sequences  Olympic  Subsequent c a l l s were pure  w h i s t l e s of a medium l e n q t h as d e s c r i b e d by Barash  to  in  prolonged  seemed t o i n d i c a t e a s t a t e of  92  Table  XI. A comparison of vocalizations within the Marmota c a l i g a t a Group  Call Characteristics  M. vancouverensis  M. caligata  C a l l Name Duration (s) Interval (s) Freguency (Hz)  long  lonq call*< »'> .56-.75<i 3> 13.6-16<»*>  C a l l flame  medium  whistle* 0.57 17.9 2910  whistle*  0.31 6.5 2910  Duration (s) Interval (s) Freguency (Hz)  C a l l Name Duration (s) Interval (s) Freguency (Hz)  short whistle* 0. 22 2.9 2910  C a l l Naire Duration (s) Interval (s) Freguency (Hz)  rapid chirps < « 0. 22 < « 2.9 ?  C a l l Name Duration (s) Interval (s) Freguency (Hz)  keeaw* 0.29 3.8 1900-1100  * <» > <*> <3> <•> <s>  M.  Cl21£U§  ( S >  2  280 0-3200<123)  descending and lcnq c a l l ascending c a l l * * ' ) 0.3{to 0.5)<i> 0.39 3<i > >5 3500< » > 2700  alarm chiip*< > 0. 1 " ' 1.3<»> 2500-3200<»3>  medium c a l l * 0.2 1-3 2700  13  accelerating c h i r p < 3 > short c a l l * 0. 1-var iatle< i > . 0.095 .05-var iable<»' 0.36 2500-3200<i> 2700 1  queeuck whistle*•> 0.3<»> 1<*> 2000-1500<»>  c a l l s used in c o r r e l a t i o n analysis cn paqe 93 Tauluian 1975 Warinq 1966 P a t t i e 1967 Gray 1967 Barash 1973b  93  The  mean i n t e r v a l between  M« c a l i g a t a ,  whistles  {r=0.91i*  with  may  be  the  n=9 p<0.01. Table X I ) .  r e s u l t from the e f f e c t o f v o c a l i z a t i o n s time  M.  vancouverensis.  and M. olympus f o l l o w s t h e same p a t t e r n . , The c a l l  i n t e r v a l was p o s i t i v e l y c o r r e l a t e d whistle  of  reguired  to  duration  of the  T h i s r e l a t i o n s h i p may on  respiration.  More  recoup expended oxygen a f t e r a long  w h i s t l e than a f t e r a s h o r t one.  M. f l a y i v e n t r i s v o c a l i z a t i o n s M. c a l i g a t a  group  in  that the c a l l  does not vary i n d u r a t i o n 1966).  M. f l a v i y e n t r i s  observation) (Gray 1967). M. olympus  when gives  other  the an  This c a l l (Barash  is  a  from  those  o f the  i s very s h o r t , 0.037 s and interval  changes  accelerando  chirp  that i s s i m i l a r to a c a l l  (Waring (personal  d e s c r i b e d f o r J|. c a l i g a t a  was not heard from M. vancouverensis 1973b).  v o c a l i z a t i o n s of H. monax. the  differ  I»loyd  (1972)  One i s a " s h o r t simple  or  described  two  whistle"  and  "two-part w h i s t l e c o n s i s t i n g of a s i n g l e high  i n t e n s i t y s h r i e k f o l l o w e d by a l e s s i n t e n s e warble."  The simple  w h i s t l e may be q u i t e s i m i l a r t o the w h i s t l e s given by s p e c i e s i n the M. c a l i g a t a group, but no sonogram was presented. part  call  The  two-  does not resemble any c a l l s so f a r d e s r c i b e d f o r any  other marmot s p e c i e s . ,  94  a l t r u i s m and Marmot Alarm C a l l i n g  a l t r u i s t i c behaviour benefits  another  can  organism,  be  defined  not  closely  d e t r i m e n t a l t o t h e organism performing and  detriment  are  defined  the  as  behaviour  that  r e l a t e d , while  being  behaviour.  Benefit  i n terms of the c o n t r i b u t i o n t o an  animal's f i t n e s s where f i t n e s s i s measured by the p r o p o r t i o n an  animal's  1974).  genes l e f t  i n the the p o p u l a t i o n gene pool  Since a marmot g i v i n g an  attention  of a predator  alarm  and thereby  call  may  subject i t s e l f  r i s k of p r e d a t i o n than i f i t had remained s i l e n t , calling  appears  to  be  altruistic.  v i r t u a l l y unknown (Pianka difficult  of  (Pianka  attract  the  to a g r e a t e r marmot  alarm  However, t r u e a l t r u i s m i s  1974) and i f present  would  be  very  t o account f o r by n a t u r a l s e l e c t i o n .  There a r e t h r e e ways t o account f o r the e v o l u t i o n of marmot alarm  calling.  selection  is  individual  The the  that  first  evolution  favour  explanation of  characteristics  if  to  that  i t i n v o l v e d some r i s k t o the c a l l e r and  considered  some  unrelated i n d i v i d u a l s .  an important  selective  alarm c a l l i n g i n b i r d s (Hamilton 1973).  The  force  i f it  an  Kin  calling  even  incidentally  Kin s e l e c t i o n has been in  the  evolution  of  1964, Maynard Smith 1965, Emlen  s o c i a l o r g a n i z a t i o n o f Vancouver I s l a n d marmots i s  such t h a t the i n d i v i d u a l s r e c e i v i n g the alarm closely  within  individual.  could account f o r t h e e v o l u t i o n of alarm  benefitted  Kin  the s u r v i v a l o f i t s c l o s e r e l a t i v e s but  that are not n e c e s s a r i l y b e n e f i c i a l selection  i s kin selection.  related  to  the  caller.  Thus,  are l i k e l y  kin  to  be  s e l e c t i o n could  95  account f o r t h e e x i s t e n c e o f alarm c a l l i n g  in  Vancouver  Island  marmots.  Secondly, is  alarm  calling  may be s e l e c t e d f o r because there  a d i r e c t b e n e f i t t o t h e c a l l e r a s s o c i a t e d with c a l l i n g .  individual  could  benefit  from  predator  (Maynard Smith  marmots  so as t o make t h e c a l l e r  individuals  (Charnov  manipulating burrow.  1965)  calling or  marmots  The r e a c t i n g  by  i f i t confused  i f i t "manipulated"  the other  l e s s v u l n e r a b l e than the other  and Krebs 1975).  other  The  Marmot  stimulating  marmots thus become more  calls them  could  be  t o run to a  conspicuous  (to  me and presumably t o p r e d a t o r s as well) than t h e c a l l e r , who has a l r e a d y moved t o a burrow entrance.  As Charnov and Krebs (1975)  argue, t h e i n d i v i d u a l s r e a c t i n g t o the c a l l "use t h e i n f o r m a t i o n to  their  own b e n e f i t , but by doing so make i t p o s s i b l e f o r the  c a l l e r t o b e n e f i t even more."  Trivers account  (1971)  presented  other  arguments  f o r the e v o l u t i o n of alarm c a l l i n g  which  by d i r e c t  would  selection  for  the i n d i v i d u a l c a l l e r . , He argues t h a t i t i s disadvantageous  for  an i n d i v i d u a l t o have a predator k i l l  because  a  nearby  conspecific  the predator may then be more l i k e l y t o k i l l  near f u t u r e .  T h i s c o u l d occur i f the predator was  him i n the  more  likely  to (1)  develop  a  search  image  (Emlen 1973) f o r t h a t prey  species, (2) l e a r n the h a b i t s o f t h e prey s p e c i e s and p e r f e c t i t s hunting techniques on i t , and  96  ,  (3) f r e q u e n t the h a b i t a t of the prey s p e c i e s .  G i v i n g alarm  calls  specializing  on  favouring  the  thus  the  tends  to  caller's  individual  prevent  species  caller  and  even  There calling.  and Krebs  may It  conspecifics  be  be  around  direct  a  mate  for  r e g a r d l e s s o f how  or  l o c a l i t y , thereby callers  neighbours  advantage  advantageous  The presence of c o n s p e c i f i c s may finding  would  be  are  (Trivers  an  closely  t o marmot alarm animal  winter  have  r e l a t e d they are.  be advantageous  increasing  to  with r e s p e c t t o  survival  due  to  r e d u c t i o n of heat l o s s when animals h i b e r n a t e i n a group. there  from  1975).  another  may  predator  though  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 1971, Charnov  a  selection  for  alarm  calling  d e r i v e d from having o t h e r animals around  exceeds  the Thus,  i f the b e n e f i t the  cost  of  calling.  Lastly,  alarm  s e l e c t i o n , which group  as  1962).  a  calling  could  i s selection  whole  rather  be  brought  favouring than  the  survival  the i n d i v i d u a l  Group s e l e c t i o n could account  for  a  about by of  group the  (Wynne-Edwards  truly  altruistic  t r a i t to e v o l v e ; however, i f group s e l e c t i o n e x i s t s a t a l l i t i s rare  (Lewontin  (Sterns 1976) i solution  1970), l e s s e f f i c i e n t and  (direct  calling) exists  i t or  should  not  than i n d i v i d u a l  be  invoked  if  selection a  simpler  k i n s e l e c t i o n i n the case of marmot alarm  ( H i l l i a m s 1966) .  The above arguments  were  presented  to  account  for  the  97  apparent  cost  ( l o s s of f i t n e s s ) a s s o c i a t e d  However, no i n f o r m a t i o n  e x i s t s f o r any  net c o s t i n v o l v e d with alarm c a l l i n g °f Marmota s p e c i e s do caller to  is  themselves  observed,;  the  (1975) saw  eight  species.  Not  risk. before  c a l l e r was  of  calling.  that there i s any  (Brown 1975).  Observations  that  the  alarm  Vancouver I s l a n d marmots appear whistling.  already  instances  one  species  not support the s u g e s t i o n  i n c u r r i n g any  protect  with alarm  In  all  cases  at a burrow entrance.  predation  of these o c c a s i o n s  was  on  various  I  Barash Marmota  preceded by alarm  calls  from the v i c t i m .  From  the  above arguments I conclude that there  altruism associated  with marmot alarm c a l l i n g . ,  c o s t at a l l a s s o c i a t e d  i s no  true  I f there i s  any  with c a l l i n g , b e n e f i t s from d i r e c t or k i n  s e l e c t i o n would be s t r o n g enough t o s e l e c t f o r alarm  The  E v o l u t i o n of Whistle  Marler  (1955,  Structure  1957)  was  p h y s i c a l p r o p e r t i e s of avian their  function.  convergently appear  He  evolved  u s i n g , barn  ears.  f i r s t t o p o i n t out how  alarm c a l l s  observed  that  some  properties of d i f f e r e n t  owls  locate  sound  could  be  bird  related species  (3?y_to.  alba) , His  a  Konishi sound  the to have  alarm c a l l s that have c h a r a c t e r i s t i c s  l o c a t i n g i t s prey by sound. owls  the  t o make them d i f f i c u l t t o l o c a t e .  the l o c a t a b i l i t y  calling.  that  (1973) t e s t e d  characteristics  s p e c i e s that i s adapted f o r  results  indicated  that  barn  by comparing the i n t e n s i t y of sound between  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 n o i s e s are e a s i e r t o  l o c a t e than pure tones because they c o n s i s t o f many f r e q u e n c i e s , each o f which can be used f o r b i n a u r a l Locatability freguency located than  by  this  method  (at l e a s t between 3 narrow  any  of  bandwidth  bandwidth the  being  other  the  is and  pure  also 9  comparisons.  directly  kHz).,  related  Thus,  to  barn  owls  tones of 3 kHz l e s s a c c u r a t e l y  sound  primary  intensity  characteristics characteristic  tested  with  determining  the  l o c a t a b i l i t y o f a sound.  Vancouver I s l a n d marmot alarm  whistles  have  the  precise  characteristics  of the most d i f f i c u l t - t o - l o c a t e sound t e s t e d by  Konishi  Narrow bandwidth alarm c a l l s are a l s o found  (1973).  many other (but by no means a l l ) s p e c i e s o f medium s i z e , mammals  (Table  XII).  A  diurnal  d i f f i c u l t - t o - l o c a t e alarm c a l l  reduce the r i s k o f p r e d a t i o n  predation  pressure c o u l d s e l e c t f o r t h e e v o l u t i o n of such alarm  calls  c h a r a t e r i s t i c s i n a l l o f the of  the  birds  narrow bandwidths. remarkably  by  convergence  in  both  birds  caller.  listed.  The  for and  that  there  fitness  of  the  same  mammals.  individuals giving d i f f i c u l t  c a l l s r e l a t i v e t o no c a l l .  Thus, s e l e c t i o n  characteristics  enter  altruistic callers) .  does  nature  of  not  marmot  alarm  has  into  alarm  been  alarm  call  Selection  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 i n d i c a t e anything the  Thus  Marler (1955, 1957) a l l had  I t appears t h e r e f o r e  strong  characteristics  recorded  species  the  would  presumably  call  to  in  for about  o r easy t o l o c a t e  for specific  the calling  discussion  call o f the  (non-callers  vs  99  The  E v o l u t i o n of Whistle F u n c t i o n  M*  vancouverensis  alarm c a l l s not only i n d i c a t e t h a t there  i s a predator around but a l s o what type of predator i s The  presence  of  shown f o r any  situation  other  specific  marmot  alarm  species.,  present.  c a l l s has not been  Such  alarm  calls  r e l a t i v e l y common among e c o l o g i c a l l y r e l a t e d s p e c i e s . terrestrial  predators  f o u r s p e c i e s of 1977,  ground  Fitch  1948;  •=>. SSSJSiatus, mountain  viscachas  Waring  1973b),  M.  1971;  peruanum,  flaviventris  (Armitage  Waring 1966), and  appear t o have s i t u a t i o n s p e c i f i c  If  situation  specific  should e x h i b i t a eight two,  specific  information  species  then  have  doqs  each c a l l .  (King 1955)  in  difficulty  in detecting  example,  the  other  Turner  1973),  1948),  fl.  and (Gray  King  (Barash  OIZIJJUS  1962  and  personal 1967)  do not  calls.  calls  receivers appropriate  are  communicating  of t h i s i n f o r m a t i o n the  s i t u a t i o n s p e c i f i c alarm c a l l s ,  only  (Turner  1973)  were observed  I t h i n k t h a t the l a c k of  responces  1966;  Of  B e l d i n g ' s ground s q u i r r e l  prairie  For  the  Balph  ludovicjanus;  M. c a l i g a t a  alarm  biologically that  1977).  et a l  Pearson  (Cynomys  et a l  and  Owings  and  beldingi,  p r a i r i e dogs Smith  Balph  S.  Avian  alarm c a l l s i n at l e a s t  (S. b e e c h e x i ,  S. armatus,  (Laqjdjum  1970,  communication.  specific  sguirrels  Melchior  perhaps b l a c k - t a i l e d 1955,  evoke  are  six small  H. vancouverensis  response.  and  to r e a c t d i f f e r e n t l y  observations  species  black-tailed  is  differences  of  probably between  to  specific due  t o the  responses.  responds to both long and  short  100  w h i s t l e s by running of  to a burrow and l o o k i n g about f o r the  the d i s t u r b a n c e .  I t would be advantageous i n t h i s  i f w h i s t l e length c o u l d provide i n f o r m a t i o n on looking  f o r a predator,  t h i s was  the only v a r i a t i o n i n the  difficult  situation  where  to  begin  e i t h e r i n the a i r or on the ground. response,it  would  be  If very  to d e t e c t or t o t e s t f o r .  Aerial  p r e d a t o r s probably  d i f f i c u l t - t o - l o c a t e alarm both  cause  situation  calls.  specific  and  (Table X I I ) , avian predators locatable  than  predators.  the  Avian  e x e r t the g r e a t e s t s e l e c t i o n f o r In a l l f i v e s p e c i e s t h a t difficult-to-locate  evoked alarm  corresponding predators  difficult-to-locate  alarm  may  call  alarm  c a l l s that  call  evoked by  select  more  that  calls less  terrestrial  strongly  characteristics  range and speed of a t t a c k i s g r e a t e r than  were  have  for  because t h e i r of  terrestrial  predators.  .The  presence  or  s i t u a t i o n s p e c i f i c alarm predation that  a  pressures prey  absence  of  c a l l s i s probably  calls.  selected  for  species  possesses.,  Situation when  specific  there  is  As avian  chances  different  of  predators.  survival  for  alarm  selective  t e r r e s t r i a l and avian predators and their  a  function  and  of  the  and the v a r i e t y of p o s s i b l e escape responses  i n c r e a s e s there i s i n c r e a s e d s e l e c t i o n alarm  difficult-to-locate  predation  difficult-to-locate calls pressure  will  only  from  i n d i v i d u a l prey can  by responding  pressure  be  both  increase  d i f f e r e n t l y to  these  101  Table X I I . A l i s t bandwidth alarm  of mammalian calls  species  that  have  narrow  Referenee  Species  M* Xancouverensis< 2> M..olympus M. c a l i g a t a M. fonax Spermophilus armatus<z > S« £ i c h a £ d s o n i i S. f r a n k l i n i i S. nndulatus< > feeechey.i<2> rock hyraxes: Hyracoidea i'SflU-iiil peruanum< 2> 2  t a g i d i a i boxi Octodon degus Chinchilla laniqer Pagyomys fiilorides P o i i c h o t i s pataqonum IJeSiolagus s a l i n i g o l a ? § c t i n a t o r sp. M j c r o c a y i a spp. C a v i a spp.  t h i s study Barash 1973b Waring 1966 L l o y d 1-972 <»•> Balph and Balph 1966 B a n f i e l d 1974<*> B a n f i e l d 1974<»> Melchior 1971 Owings e t a l 1977 Mathews 1971<i> Pearson 1948 and Eisenberg 1974 Rowlands 1974<»> E i s e n b e r g 1974< »•> Eisenberg 1974< »> Eisenberg 1974< * y Eisenberg 1974< i > Eisenberg 1974< * > George 1974<»> Rood 197 2: 16<»> Rood 197 2: 17<D  <ir - No sonograms were presented but t h e v e r b a l d e s c r i p t i o n appears t o be a p p r o p r i a t e , and a narrow bandwidth sound i s r e l a t i v e l y easy to d e s c r i b e . ( z ) - Alarm c a l l s a r e a l s o p r e d a t o r - s p e c i f i c .  102  In  addition  whistles  may  predator  to  warning  reduce  other  predation  i s aware t h a t i t has  marmots  seek  cover,  by i n h i b i t i n g a t t a c k s .  I f the  been detected  otherwise  auspicious  probability  of c a p t u r i n g a l e r t e d prey.  (1975)  noted  attacks. birds  that  attack  because  it of  may  Warning c a l l s may  Alcock  a l s o f u n c t i o n as  abandon  the  overall  (1975) and  alarm c a l l s or s i g n a l s may  by d i s c o u r a g i n g a predator  (Wilson  to  inhibit  mobbing  from remaining  an low  Brown  predator calls  in  i n the v i c i n i t y  1975). ,.  INTERSPECIFIC COMPARISONS  OF  MARMOT SOCIAL BEHAVIOOS  A Test of Barash*s Hypothesis  ;  In order to t e s t t h e data obtained  Barash*s  hypothesis  in  this  of marmot s o c i a l i t y , i t was  to o b t a i n e x p l i c i t p r e d i c t i o n s p e r t a i n i n g marmots.  The  parameter necessary  estimate  of  the  length  of  to  study first  against necessary  Vancouver  Island  t o make such p r e d i c t i o n s i s an  the v e g e t a t i v e 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 of  (Barash  1973b).  frost-free  (Table I I ) .  days  The  of  was  115  days i n 1973  and  average number of f r o s t - f r e e  s u b a l p i n e parkland number  On the Haley Lake study  h a b i t a t i s 114  frost-free  days  was  but  in  equivalent  1973  persisted  weeks a f t e r the l a s t f r o s t .  v e g e t a t i v e growing season t o about  the  number  135 days i n days  in  1974  121  large  days.  to the amounts  1974  typical  days (Brooke et a l 1970).  growing season i n f o r two  area  The  vegetative of  snow  T h i s reduced the Snow  typically  103  persists  for  environments  about  3  weeks a f t e r t h e l a s t f r o s t  {Brooke et a l 1970), thus averaqe growing season i n  subalpine environments i s about 93 days. growing  season  of  40  intermediate  The  93  to  150  to  70  days,  growing  days  therefore  day  M. olympus e x p e r i e n c e s a short growing H. fla.yj.vejntr.is  experiences  (Barash  predict  an  season o f 70 to 100 days, and H. monax i n  c e n t r a l Pennsylvania e x p e r i e n c e s a very l o n g growing about  121  i s r e l a t i v e l y l o n g compared to t h a t experienced  by other marmot s p e c i e s . season  i n subalpine  that  1974a). the  Barash's  social  season  hypothesis  behaviour  of  would  and  social  o r g a n i z a t i o n o f H. vancouverensis should be very s i m i l a r t o t h a t of  M.,flaviventris.  That  1) be s o c i a l l y i n t o l e r a n t greeting  rate,  i s , Vancouver I s l a n d marmots should as  indexed  by  having  a  very  low  2) be moderately a g g r e s s i v e , 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  distinct  home  ranges,  and  4) grow  q u i c k l y and d i s p e r s e at an e a r l y age, probably as y e a r l i n g s .  Individuals social  group  would  individuals greeting perhaps Barash  belonging  belonging  presumably  1973a,  to  a  reinforces  1973b,  h i g h l y i n t e g r a t e d and to  less  greet  more  cohesive  individual  been p o s t u l a t e d and  1974b;  Steiner  stable  often  society  than because  recognition  1975).  (King  and 1955;  Individual  on the b a s i s of scent has been demonstrated i n many  mammalian s p e c i e s  mountain  likely  a  a l s o f u n c t i o n s as a method o f scent s h a r i n g  recognition  spp.),  be  to  (Halpin for  prairie  sheep  1974, Shorey  marmots, doqs  ground  1976).  Scent s h a r i n g  squirrels  (Cynomys spp., S t e i n e r  (Ovis spp., G e i s t  1971).  has  (Spermophilus 1975)  and f o r  104  A comparison o f Vancouver I s l a n d marmot g r e e t i n g r a t e s with those pf other marmots r e v e a l s t h a t Vancouver I s l a n d marmots a r e among the most s o c i a l o f  a l l marmot  species  {Fig  g r e e t i n g r a t e of M. vancouverensis i s much higher estimates  of  the  rate  The  greeting  prediction  rate  of  marmots  (Barash  based  Thus  M. marm.ota  the  are  on  Barash's  of  of  somewhat  both  are  greetings  suspect,  at  pattern  that  (1976b)  significantly  and  least  1976b, yellow-  1973a, F i g 3 ) ; however, Barash  rates  This  (Barash  than that  s t a t e s t h a t the g r e e t i n g r a t e s of M. maoiot§ lower.  than a l l three  M. marmota  T a b l e s I I S IV) appears to be much higher bellied  The  at which M. f , l a y j y e n t r i s g r e e t s .  r e s u l t i s i n c o n s i s t e n t with the hypothesis.  26).  chases  the  way  I  in am  i n t e r p r e t i n g them.  The chasing  only  behaviour  states  rate that  significantly  was  (Barash 1973b) .  the  freguency  between  the  of  assumed  these I  chasing  chasing two  could rate.  s o c i a l l y t o l e r a n t species.  did  species. compare  the  same  in  Thus  differ the  only  most  marmot  with  less  social  chases  more  freguently  Chasing r a t e v a r i e d l e s s among  marmot s p e c i e s than d i d g r e e t i n g r a t e s ,  with  having  ( F i g 27).  a r e l a t i v e l y low r a t e o f chasing  r a t i o of g r e e t i n g s t o chases i s  not  among  Species  t o l e r a n c e would l i k e l y p a r t i c i p a t e than  I  present  f o r M. olympus as M. marmota s i n c e Barash (1976b)  a g o n i s t i c behaviour t h a t species  g u a n t i f i e d was  (Barash 1973a, 1974b, 1976b) and he d i d not even  t h a t data f o r H. olympus chasing  Barash  probably  the  H. vancouverensis  most  However, the meaningful  105  Figure 26. & comparison 1 2 3 4 5a 5b 5c 6  _  -  of g r e e t i n g r a t e s among marmot  species  Marmota vancouverensis ( t h i s study) - I~ olympus (Barash 1973b) ~ M. c a l i g a t a (Barash 1976b) M. marmota (Barash 1974b) - H. f l a v i v e n t r i s (Armitage 1974, 1976a) - 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) - 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) - M. monax (Bronson 1964)  O  106  paramater  t o compare  reasons f o r t h i s .  among  marmot  First, socially  species.  There are three  intolerant species  would  be  expected t o have a g o n i s t i c a c t s making up a h i g h e r p r o p o r t i o n o f their  social  interactions.  Second,  a comparison o f the r a t e  r a t i o s should overcome the d i f f e r e n c e s methods  used  to  calculate  comparison of r a t e r a t i o s colony  size.  When  marmot  Barash's  interaction  i s independent  the r a t e s  compared, Vancouver I s l a n d social  species  among  of  marmots  authors rates.  of  greeting appeared  (Fig 28), a  Third,  any to to  i n the  effects chases  be  third  Vancouver  prediction  Island  of  most  result inconsistent  with  hypothesis  is  T h i s was c l e a r l y not the case. home ranges  w i t h i n a colony or, i n t h e case o f colony one, w i t h i n a colony.  of  a  Complete home range o v e r l a p  was  a l s o found i n M. olympus 1974b).  that  marmots should have i n d i v i d u a l t e r r i t o r i e s or  Vancouver I s l a n d marmots have completely o v e r l a p p i n g  (Barash  section  among c o l o n y members  1973b)  and  M. c a l i g a t a  Y e l l o w - b e l l i e d marmots a r e grouped i n t o harems  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  1962,  were  the  Barash's  r e l a t i v e l y d i s t i n c t home ranges.  but  of  hypothesis.,  The  (Barash  a  feeding  areas  (armitage  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  final  Vancouver disperse  Island  prediction marmots  of  Barash's  should  hypothesis  slow,  as  yearlings  that  grow and mature g u i c k l y , and  a t an e a r l y age, presumably as y e a r l i n g s .  relatively  is  of both  sexes,  Growth  was  two-year-old  107  F i g u r e 27. A comparison 1  ~ ~ 4 ~ 5b 5c 6 2  3  !  5  : !  ;  ':;  a  of c h a s i n g r a t e s among marmot s p e c i e s  Marmota vancouverensis {this study) !!• olxmjDus {see t e x t ) M. c a l i g a t a (Barash 1976b) M. marmota (Barash 1974b) M« f l a v i v e n t r i s i Armitage 1974 Barash 1976a) M. f l a v i v e n t r i s (high e l e v a t i o n , Barash 197 3a) 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) M. monax (Bronson 1964) f  05  o  JZ I  c  CTJ  tf) - 0 . 0 h C O  '•+-» U  (?  (TJ O)  c (TJ JZ  U Q  -1.0 h  -2.0 h  108  F i g u r e 28.  A comparison o f t h e among marmot s p e c i e s  1 2 3 4 5a 5b 5c 6  -  ratio  of  greetings to  chases  Marmota v a n c o u v e r e n s i s ( t h i s s t u d y ) |f. o i l ! p u s ( B a r a s h 1973b) M. c a l i g a t a ( B a r a s h 1976b) a. marmot.a ( B a r a s h 1974b) M. f l a v i v e n t r i s ( A r m i t a g e 1974, 1976a) M. f l a v i v e n t r i s - ( h i g h e l e v a t i o n , B a r a s h 1973a) M. f l a v i v e n t r i s (medium e l e v a t i o n , B a r a s h 1973a) M. monax ( B r o n s o n 1964)  108a  CD H—'  03  cr  1.0  U)  c ro  U -*-»  I  c  CO if)  c  CTj  o 2  O)  CD  c V  0.0  -1.0  -20  CD  o Q  All Age and Sex Classes Combined  109  females,  and  perhaps  distinguishable  from  also  adults  two-year-old by  weight  growth r a t e can be obtained by comparing from  their f i r s t  adults  at  hibernation  that  marmots grew  time  slower  ( F i g 4).  year  than  both  were  An index of  marmots a f t e r emergence  (yearlings)  of  males,  with  (Table XIII) .  the  weight  of  Vancouver I s l a n d  M. f l a v i v e n t r i s  and  M. monax  (Table X I I I ) .  Vancouver  Island  marmots  appear t o have delayed m a t u r i t y  e q u i v a l e n t t o that of M. olympus. that old  I was  Two  Vancouver I s l a n d  sure had d i s p e r s e d (males #4 and #17)  were two  or o l d e r , while n e i t h e r of the two two-year-old  my . study  areas  produced  litters..  had l i t t e r s  either  colony,  presence  on  the  t r a p p i n g or the l a t e s p r i n g i n 1974  years  females  on  However, none of the a d u l t  females on c o l o n i e s one or two my  marmots  my  were  i n 1974  either.  disturbance responsible  If  through for  the  a d u l t females not breeding then the same f a c t o r s would be a c t i n g on  both  two-year-old and a d u l t females. , I f any of these three  reasons were r e s p o n s i b l e f o r a d u l t females not breeding  then  I  would have no b a s i s f o r p o s t u l a t i n g t h a t two*year-old females do not  breed  because  they  are  f a c t o r s can be d i s c o u n t e d : 1) 1973  immature.  However, these three  Three females produced  when I observed but d i d not t r a p .  T h e r e f o r e my  the colony was not s u f f i c i e n t t o i n h i b i t b r e e d i n g . one  adult  she s t i l l  female on the Ski Club colony produced  a litter,  in  presence on 2) I trapped  (colony 5) i n 1974  and  thus t r a p p i n g does not appear t o be  s u f f i c i e n t t o i n h i b i t breeding. i n t o the s p r i n g of 1974  litters  3) The p e r s i s t e n c e of snow l a t e  was not s u f f i c i e n t  to  inhibit  breeding  110  t a b l e XIII. a comparison of the ? y e a r l i n g marmots  relative  Y e a r l i n g / a d u l t Height i n the Spring (%)  Species  M... M; H. M. M." 1• H« M.  calicjata vancouverensis olympus flaviventris flaviventris flaviventris flaviventris monax  i z  high e l e v a t i o n c o l o n y medium e l e v a t i o n colony  1  2  18 44 30 50 56 75 70 65  growth  rates  Reference  Barash 1976a t h i s study Barash 1973b J . Donaldson i n prep armitage e t a l 1976 Barash 1973a Barash 1973a Snyder et a l 1961  of  111  since  breeding  occurred  on  other c o l o n i e s  (e.g. S k i Club and  B u t t l e r Peak) even though the snow c o n d i t i o n s were very in  a l l areas.  The most l i k e l y  similar  e x p l a n a t i o n f o r the f a i l u r e o f  a d u l t females on c o l o n i e s one and two t o breed i n 1974, Vancouver  Island  marmot  females  may  is  only breed i n a l t e r n a t e  y e a r s , a r e p r o d u c t i v e s t r a t e g y s i m i l a r to M. olympus. breeding  could  account  because jthree l i t t e r s  but  one  the  absence  had been born the  marmots were not tagged that produced l i t t e r s  for  i n 1973,  that  of  Biennial  litters in  previous  year.  1974 Since  I cannot be sure that a l l t h r e e  i n 1973 were the same ones present i n  distinctively  marked  individual  1974  was present i n both  years.  ,None o f the four p r e d i c t i o n s of consistent Vancouver  with  observed  data  on  hypothesis  the  Inconsistent  species  results  c o n s t r u c t a new  with  such  the  as  marmot  species  should  was  trying  growing  season  why  some  others  to  reach  M. f l a v i v e n t r i s ,  adult  M. monax,  or  vegetative  to The  growing  f o r i n d i c a t i n g the degree of s o c i a l  guestion t h a t Barash was  season.  of the e x i s t i n g one.  to r e j e c t i s , that  season i s a u s e f u l parameter a  growing  these lead one t o e i t h e r t r y to  one or more o f the assumptions  tolerance  shortest  resembled  h y p o t h e s i s , having r e j e c t e d the o l d one,  most obvious assumption  were  M. vancouverensis.  I s l a n d marmots i n a l l i n s t a n c e s most c l o s e l y  M. olympus,  modify  the  Barash*s  to  answer  The when  fundamental considering  marmot s p e c i e s take l o n g e r than  size. and  exhibit.  He  M. olympus  found the  growing season c o r r e l a t e d with the l e n g t h of time  that  among  length o f the marmots  took  112  to  mature.  Anderson  However,  et a l  Vancouver  the  (1974)  island  are  when  with  93-121  day  more  marmots  this  growing  marmots mature one year e a r l i e r  trend.  (breed)  until  season  while  when the growing et a l  1974).  data are c o n s i d e r e d , v e g e t a t i v e growing  season  does not appear t o guickly  inconsistent  a  season i s only 60-100 days Thus  o f t h i s study and those of  marmots do not appear t o mature  t h e i r f o u r t h summer with yellow-bellied  results  be  a  (Barash 1974a, Anderson  meaningful  reach maturity.  parameter  Anderson  affecting  how  et-al-(1974) suggest  that the l e n g t h of time that marmots grow throughout the  summer  (the marmot growing season), would be a more a p p r o p r i a t e measure of  environmental  severity.  However, the l e n g t h of the marmot  growing season does not c o r r e l a t e any b e t t e r with the time taken t o reach maturity  than  does  the  vegetative  growing  season.  . . . . . . i,  Olympic  marmots  gain  while y e l l o w - b e l l i e d days  weight  f o r up t o 120 days  (Barash 1973b)  marmots a t high e l e v a t i o n s grow f o r o n l y 96  (Anderson e t a l 1974) and mature one year e a r l i e r .  the marmot growing season nor the v e g e t a t i v e growing correlated  with  the  reach a d u l t s i z e i s independent  Neither  season  are  age o f maturity because the time taken t o  determined  factors;  by  the  combination  of  three  1) the r a t e t h a t marmots put on weight i n  terms of grams per day,  2) the  continue  to gain weight  (the marmot growing season), and 3) the  absolute  weight  considered.  of  These  an three  adult  number  marmot  factors  of  of  varied  days  the  that  marmots  species  being  among s p e c i e s but I  could not determine any c o n s i s t e n t u n i f y i n g t r e n d s among them.  The most obvious way t o  determine  how  long  i t takes  a  113  marmot  to  directly. the  reach  adult  of  the g r e e t i n g  marmots 1} at the end  2)  of the f a l l  the  logarithm  to  measure  negative rate and  of t h e i r second  of  the  weight  a d u l t s at t h a t time of year  the  (r=0.95,  between  weight of immature expressed  <r=0.86, n=6,  of  hibernation  the growth r a t e  correlation  summer,  weight of an a d u l t  emergence from t h e i r f i r s t  XIII)..  is  There i s a s i g n i f i c a n t  logarithm  percent  size  yearlings  as  p=0.03) and after spring  r e l a t i v e to the weight of n=7,  p=0.0 01,  and  Table  Thus, by measuring the growth r a t e d i r e c t l y r a t h e r  estimating relative  i t from the l e n g t h of the v e g e t a t i v e social  tolerance  can  be  a  growing  predicted  for  than  season,  a l l marmot  species.  The  reason that Vancouver I s l a n d marmots take a  t o mature i s not  that they grow s l o w l y  long  time  i n terms of grams per  day  or because they only gain weight f o r a short p e r i o d each summer, for  they  are i n t e r m e d i a t e  both of these parameters. time  to  size.  A  increases  among marmot s p e c i e s with r e s p e c t Vancouver I s l a n d marmots take a  l a r g e a d u l t body  large  for  body  size  may  be  selected  the chance of s u c c e s s f u l d i s p e r s a l .  successful  However,  the  dispersal  to  probability  i n c r e a s e s with body s i z e .  new of  successful  T h i s would occur  is  the p r o b a b i l i t y probably  dispersal if  larger  o f time without food and  s h e l t e r than could  adults  causes  low.  probably  f a s t e r , and s u r v i v e f o r a longer  from  it  Vancouver I s l a n d  h a b i t a t and habitats  because  could t r a v e l f u r t h e r and  aggression  long  mature because they have a r e l a t i v e l y  marmots l i v e i n i s l a n d s of subalpine of  to  animals period  s m a l l e r animals.  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Sound communication of t h e y e l l o w - b e l l i e d 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 , whitet a i l e d and Gunnison«s p r a i r i e dogs. Am. M i d i . Nat. 83; 167185. Williams, G. C. 1966. Adaptation and Natural Selection. P r i n c e t o n U n i v e r s i t y P r e s s , P r i n c e t o n , N.J. 307 p. Wilson, E. 0. 1975. S o c i o b i o l o g y : The New S y n t h e s i s . U n i v e r s i t y P r e s s , Cambridge, Mass. 697 p.  Harvard  Wright, H. E., and D. G. Frey, ed. 1965., The Quaternary o f United S t a t e s . P r i n c e t o n U n i v e r s i t y P r e s s , P r i n c e t o n , N.J. 922 p. Wynne-Edwards, V. C. 1962. Animal D i s p e r s i o n i n R e l a t i o n S o c i a l Behaviour. O l i v e r and Boyd, London. 653 p.  to  124  Yeaton, R. I. 1972. S o c i a l behaviour and s o c i a l o r g a n i z a t i o n 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 P l a n t Species Found on the Haley Lake Study Area I n d i c a t i n g Those Species Known t o be Eaten by Vancouver I s l a n d Marmots  Family and S c i e n t i f i c Same  Common Name  Selaginellaceae Selaginella wallacei  selaginella  Polypodiaceae Adiantutn pedatum Cryptogramma c r i s p a Pply_stichum muni turn Pteridium aquilinum  maidenhair f e r n parsley fern sword f e r n bracken f e r n  Cupressaceae Chamaecyparis n o o t k a t e n s i s J u n i p e r i s communis  yellow cedar dwarf juniper  Pinaceae Abies a m a b i l i s Abies lasiocarpa Pinus conto r t a Pinus monticpla Tsuga h e t e r o p h y l l a Tsuga. mertensiana Pseudotsuga m e n z i e s i i  amabilis f i r alpine f i r lodgepole pine western white pine western hemlock mountain hemlock Douglas f i r  Betulaceae Alnus s i n u a t a  Sitka  alder  Aristolochiaceae *sarum caudatum  wild  ginger  Portulacaceae Clay_tonia l a n c e o l a t a  western springbeauty  Caryophylaceae A r e n a r i a macrop_hylla Silene menziesii  b i g l e a f sandwort catchfly  Ranunculaceae Actaea rubra M M i i S a i S . formosa Delphinium sp. . Thalictrum Occidentale  banenerry columbine larkspur meadow rue  P l a n t s Eaten  fronds  0  bark°  lvs», f l w s  2  l v s , flws, fr'  3  126  Berberidaceae ASfeilus t r i g h y l l a  vanilla  Crassulaceae Sedum diyerqens  stonecrop  Saxifraqaceae . ; S a x i f rag a b r o n c h i a l i s ferruginea . .Saxifraga o c c i d e n t a l i s ' T e l l i m a grandiflorum  spotted s a x i f r a q e rusty saxifraqe western s a x i f r a g e fringecup  Grossulariaceae Ribes l a c u s t r e  swamp  Rosaceae f£a<jaria y i r g i n i a n a Luetkea p e c t i n a t a Potentilia diyersifolia Sorbus s i t c h e n s i s  b l u e l e a f strawberry partridgefoot cinquefoil S i t k a mountain ash  Leguminosae Lathyrus neyadensis Lnjjj.nus l a t i f o l i u s  sweat-pea broadleaf lupine  Celastraceae Pachistima m y r s i n i t e s  f a l s e box  Violaceae Viola glabella  yellow  Onagraceae Epilobium  a l p i n e fireweed  alpinum  Ombelliferae Heracleum lanatum Lqmatium sp. Ericaceae A r c t o s t aph y1gs u y a - u r s i Cassiope mertensiana Phyllodoce empetriform i s Ifeo^odendron a l b i f l o r u m Vat eg i n i u m c a e s g i t o s a Vaccinium d e l i c i o s u m Vaccinium membraniceum laccinium ovalifolium laSSiSiSS  spp.  leaf  gooseberry  lvs  violet  cow-parsnip biscuit-root  l v s * , flws*  lvs, f r * kinnikinnick white moss-heather lvs° r e d heather white rhododendron dwarf huckelberry b l u e - l e a f huckelberry t h i n - l e a f huckelberry o v a l - l e a f huckelberry huckelberry lvs, f r *  Polemoniaceae Phlox d i f f u s a  spreading  Labiatae ; liSSS111 v u l g a r i s  self-heal  phlox  l v s , flws*  127  Scrophulariaceae £astilleja miniata Castilleja paryiflora UiJBiiiiJS g u t t a t u s ge'dlcularis b r a c t e o s a Penstemon d a y i d s o n i i ' 2§I2SS£a wormsk-ioldii  I n d i a n paintbrush Indian paintbrush yellow monkey flower lousewort penstemon speedwell  Rubiaceae Galium boreale  northern bedstraw  Caprifoliaceae Sambucus racemosa  red  Valerianaceae Valeriana s i t c h e n s i s  elderberry  lountain v a l e r i a n  ns*, f l w s * ns, f l w s  bark lvs  Campanulaceae Campanula r o t u n d i f o l i a  bluebell  flws  Compositae Achillea millefolium Agosetis aurantiaca , Anajghalis margarjtacea Arnica l a t i f o l i a C i r s i u m edule ' Erigjeron £eregrinus Efpphyllum l a n 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  yarrow agoseris pearly everlasting broad-leaf arnica Indian t h i s t l e mountain d a i s y woolly sunflower g i a n t ragwort common dandelion  lvs°  Juncaceae Juncus drummondii  rush  Cyperaceae Carex m e r t e n s i i Carex n i g r i c a n s Carex s p e c t a b i l i s Carex spp..  sedge sedge sedge sedge  lvs°  l e a f t i p s , ns lvs,  Graminae Agrostis diegoensis Bromus s i t c h e n s i s Elymus glaucus Mellcia sub a l a t a Phleum alpinum Trisetum spicatum  bentgrass brome grass blue w i l d - r y e Alaska oniongrass mountain timothy spike trisetum  Lilaceae Alium crenulatum E'rit'hrpnium g r a n d i f l o r u m L i l l u m eg1umbi anum Smilacina racemosa Stenarithium O c c i d e n t a l e T r i l l i u m ovaturn Veratruffl y i r i d e  w i l d onion avalanche l i l y tiger l i l y f a l s e Solomon*s s e a l western stenanthium western t r i I l i u m f a l s e hellebore  lvs, lvs,  lvs  ns.  ns, flws flws*  128  Orchidaceae Habenaria sp.  bog  Lichens Cladonia s p . P e l t i g e r a apthosa Solorina crocea Vstereocanlon sp. T h a n o l i a • s p . ., Umbilicaria s p . ,  rock  * z 3 * * °  leaves flowers fruit new shoots p r e f e r r e d food r a r e l y eaten  orchid  tripe  thalli  Appendix  II. A  List of a l l Collected, Total =  Known 30  Vancouver  Island  Marmot  Specimens  that  have  been  CM  B.  S.  A.  Peak  D.  I . Walker  Collection Localities*»>  Year Collected  Collector  Swarth  '  7 ca)  1910  Mt. Douqlas King Solomon Golden Eagle  Easin Easin  3 1  1929  Eattle dt. ( l o c a t i o n unknown)  1  1930  Jordan  Oruyn Mountain colony one  P.  S.  Martin  1938  Ht.  1.  E.  Gatenly  19U0  Drink  P.  L.  Deebe  D. G. King P.  L.  Bright  J.  Csman  Heard  Water  Creek  Berkley,  British  Columbia  r  12 f> 0.  II IT  hor  Provincial  Museum,  rin Ivor::!  t y o f BrU. l;ih C o l u m b i a , nuntors 1-S8f»fi a n d » 9 2 B (n = 7) , 6 N a t i o n a l Museum o f C a n a d a , C t t a w a , n u m b e r s 1«088-m089 (n = 2) .  1(3>  B r i t i s h Columbia nunter 2898.  Provincial  Museum,  1 CJ>  B r i t i s h Columbia n u r c t e r U5U0.  Provincial  Museum,  British  Provincial  Museum,  Columbia  19U3  Mt.  Washington  1  1965  Mt.  Washington  1<*>  Oniversity of nunter 28751.  1968  Rt.  Washington  1  U n i v e r s i t y o f Montana, n u n t e r UMZM 1 3 5 2 1 .  1968  Comox  1  E r i t i s h Columbia un <ataloqued.  1<3>  E r i t i s h C o l u m b i a F i s h and W i l d l i f e Branch, Port A l b e r n i O f f i c e  1  Oniversity of B r i t i s h unca t a l o q u e d .  Columbia,  U n i v e r s i t y of B r i t i s h teaching collection.  Columbia,  1968  D. C.  Arrowsmith  University of C a l i f o r n i a , n u t t e r s 12090-12100.  N a t i o n a l Museum o f C a n a d a , O t t a w a , number 1 0 3 3 3 .  Meadows  ft. Pacey and . 1931 I. t e l . Cowan  Present Location of Specimens  Number Collected  1971  ?  Beaufort  Range  Green Mountain colony one  nunter  <3>  1(3)  <o <«> < 3> to  5021. Alaska,  Provincial  mapped c n 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 t h e c c l m c l e t e s k e l e t o n skull crly . ,, . . s k i n o n l y , may go w i t h t h e s k u l l i n t h e DEC t e a c h i n g c o l l e c t i o n ^ t  Museum,  

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