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Space use in a population of least chipmunks in the Southwest Yukon Glennie, Linda Cuffableness 1988

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SPACE USE  IN A POPULATION  OF LEAST CHIPMUNKS  IN THE SOUTHWEST YUKON  by  Linda B.Sc,  Cuffableness McGill  A THESIS SUBMITTED  Glennle  University,  1984  IN PARTIAL FULFILLMENT OF  THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE  in THE FACULTY OF GRADUATE STUDIES (Department o f Z o o l o g y ) We a c c e p t  t h i s t h e s i s as conforming required standard  to the  THE UNIVERSITY OF BRITISH COLUMBIA September 1988 ©Linda  Glennle,  1988  In presenting  this thesis in partial fulfilment  of the  requirements for an advanced  degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department  or  by his  or  her  representatives.  It  is  understood  that  copying or  publication of this thesis for financial gain shall not be allowed without my written permission.  Department of  Zoology  The University of British Columbia Vancouver, Canada  D a t  e  DE-6 (2/88)  9/9/88  i i ABSTRACT  This  thesis  chipmunks a t  d e s c r i b e s an i n v e s t i g a t i o n  Kluane  demography,  home  population.  Based  summers, mean similar  number  Chipmunks  which  range  measured  i n any p r e v i o u s l y  use.  to defend neighbours weight,  using  core  suggested  provoked  Comparing  the  encounter  encounters,  except  Grid-trapping enough t o a f f e c t  o v e r two  the previous  forest  found  four  i n the  s t a b i l i t y was  and s h r u b - l a n d t o  of the  t e l e m e t r y averaged  between s o c i a l 93.4%;  genus.  Home  4.86 ha, h i g h e r  that  interactions  dominance  r a t h e r than were such  s p a c i n g and s p a c e  c h i p m u n k s do n o t a p p e a r  Provoked  social  interactions  suggest  by a  grids  i s generally  typical  areas.  that  data  generated  i n the  p u b l i s h e d s t u d y of the genus.  telemetry  that  i s also  and b r e e d i n g c o n d i t i o n ,  Although  than  o v e r l a p averaged  exclusive  two  there over  open  the r e l a t i o n s h i p  Home r a n g e  from  I examined  patterns  although year-to-year  preferred  forest,  I examined  was l o w e r  south,  Yukon.  use  data  use i n l e a s t  on t h e s t u d y a r e a was 2 2 . 6 / g r i d ,  of animals  closed-canopy  than  habitat  numbers measured  same s p e c i e s f u r t h e r  sizes  and  southwest  on l i v e - t r a p p i n g  The p o p u l a t i o n  typical.  i n the  range  t o chipmunk  years.  Lake,  of space  was b a s e d ownership  behaviour  home r a n g e  showed  that  i n c r e a s e food  size.  of space.  was a r t i f a c t u a l .  There  animals t o  chipmunks  when h a r v e s t i n g s e a s o n a l l y a b u n d a n t d i d not  on a g e ,  g e n e r a l l y aggressive, the  frequency of r a d i o - c o l l a r e d  random model  among  or cover  avoided  food.  availability  was e v i d e n c e  that the  ill r  presence  of  invalidate and  traps affected  t r a p - b a s e d home  telemetry  significant  based  use range  estimates  differences.  distribution/ estimates. of  home  b u t n o t enough t o Comparison  range  size  of trap  yielded  no  iv  Table  of Contents  ABSTRACT Table  i i  of Contents  List  of Tables  List  of Figures  iv v i i viii  ACKNOWLEDGEMENTS  ix  CHAPTER 1:  GENERAL  CHAPTER 2:  DEMOGRAPHY,  LEAST CHIPMUNK  INTRODUCTION HABITAT USE,  AND HOME  RANGE OF THE  IN THE SOUTHWEST YUKON  INTRODUCTION METHODS  1  3 .  3  . .  4  Study area  4  Live-trapping  4  Radio-telemetry  8  Habitat  9  classification  RESULTS  .' . . .  12  Demography  12  Home Range  19  Habitat  20  preference  DISCUSSION  28  Demography Habitat  use  Home r a n g e SUMMARY  .  28 32 33 36  V  CHAPTER  SPACE  3:  USE  AND SOCIAL STRUCTURE  IN  THE LEAST  CHIPMUNK  37  INTRODUCTION  37  METHODS  39  Behavioural Encounter  observation  avoidance  39  . . . .  41  RESULTS  45  Home r a n g e  overlap  Behavioural Encounter  45  observation  . .  avoidance  53  DISCUSSION  CHAPTER  54  THE INFLUENCE  4:  HOME RANGE  51  IN  OF GRID-TRAPPING ON  SPACE USE  AND  THE LEAST CHIPMUNK  60  INTRODUCTION  60  P r e d i c t ions  61  METHODS  .  Experimental Home r a n g e Effect  of  design  . . . . .  calculations data  source  .  62 62 63  on home r a n g e  estimate  RESULTS  . . .  64 65  Effects  of  grid-trapping  Effects  of  data  source  on home r a n g e  on home r a n g e  DISCUSSION Effects  .  65 67 67  of  grid-trapping  on home r a n g e  67  vi Effects  of  d a t a s o u r c e on home r a n g e  LITERATURE CITED APPENDIX:  SOURCE  69 71  CODE  LISTING  ENCOUNTER SIMULATION MODEL  (in  BASIC)  . . . .  OF RANDOM CHIPMUNK 76  vii List  o£ Tables  Table 2.1—Summary of g r i d - t r a p p i n g changes Table 2 . 2 — H a b i t a t  8  classifications  Table 2 . 3 — P o p u l a t i o n  . .  means  10 15  Table 2.4—Mean Minimum S u r v i v a l Rates  16  Table 2.5—Minimum over-winter  17  survival  Table 2.6—Means (and ranges) of chipmunk weights Table  2.7—Summary  of  habitat  preferences  ( l n g) . .  from  trapping  data Table  2.8—Chipmunk  18  25 home  ranges  from  this  and p r e v i o u s  studies  34  Table 3.1—Mean home range o v e r l a p  46  Table  i n t e r a c t i o n to home . . . . .  52  frequency on g r i d H v s . s i m u l a t i o n  54  3.2--Mean d i s t a n c e s from  Table 3.3—Mean encounter Table 4 . l--Telemetry-based  home range e s t i m a t e s  *  65  viii List  of Figures  Fig.  2.1 —  The s t u d y  Fig.  2.2 —  Jolly-Seber population  area  6  . estimates  f o r Grids  X and  H  13  Fig.  2.3 --  Fig.  2.4  Habitat —  Observed  locations Fig.  2.5 — habitat  maps:  and  vs. vegetation  Habitat  21  ( a ) G r i d X, ( b ) G r i d H  use index  expected  number  of t e l e m e t r y 26  type foradults  and j u v e n i l e s v s . 29  type.  Fig.  3 . 1 — Flowchart  Fig.  3 . 2 —Telemetry-based  home r a n g e map o f G r i d H  47  Fig.  3 . 3 —Telemetry-based  home r a n g e map o f G r i d X  49  o f random-encounter  s i m u l a t i o n model.  43  Ix ACKNOWLEDGEMENTS  I would research their  like  to  committee,  advice  Jamie  and  various Carole  Patty  Sinclair  for  S m i t h ' s a m a z i n g l y prompt  and  Gass  Jamie  field  the  with  Mclntyre  suggestions,  programming,  and  possible.  these  would  be  locked  assistance,  Chris  Inside  Blachford Foote  C o u r t e n a y , G o r d o n Haas, Rob  field  Sian  and  his  and  Megan  Wright,  provided invaluable  f o r m a t t i n g , and help  the  in  chipmunk  computer. Susan  once  Powell  and  Watts  b r o w - b e a t i n g , and  the  Andy  of  assistance.  Peter  Without  (who  of  help  Mowat, S a l l y  proof-reading,  acknowledgements.  still  Garth  provided  constant  provided  encouragement  stage,  figures,  Tony  my  were g r e a t l y a p p r e c i a t e d .  work  wrlting-up  and  Station a l l  friendship  Lisa  I thank A l i s t a i r  and  Lee  rough d r a f t  made t h e  H a r r i s , and  writing  K r e b s and  Andrew L a w r e n c e , S h i e l a Fox,  innumerable  data  The  Williams  During  help  the  supervisor, Charles  of K l u a n e R e s e a r c h  kinds.  Williams,  Smith,  support.  t h o r o u g h comments on Inhabitants  t h a n k my  and  Ertis  saved Don  for  my  programming  life),  Robinson  for  Simon advice  for b o l s t e r i n g morale. I thank  convinced My  of  my the  field  mother  for being  u t t e r lunacy work  funded  Grants,  and  teaching  a s s i s t a n t s h i p s while  This  NSERC  was  of  grants  thesis is dedicated  to at  endlessly supportive this  while  endeavor.  by Dr.  even  Northern Krebs.  Studies I was  U.B.C.  t o Cygnus and  Zombie.  Training  supported  by  1  CHAPTER  This  study  population the  of  on  the  Since of  chipmunks  use,  habitat  have  (Heller,  included  1978).  competition habitat the This  other  study  isolation  from  Chapter  2  studies  the  in a  dominance  i n the  studied  1976; States, 1976;  and  the of  influence  of  differences in  i n the d i s t r i b u t i o n of sympatric  on  space  the  congeners.  use,  least  habitat  chipmunk  in  i t s congeners. provides  an o v e r v i e w o f demography and home r a n g e  It describes  habitat  use a r e  studies  of the genus. 3  for  patterns  among t h e s p e c i e s  zonation  dominance  o f chipmunk.  F i v e of  assessed  data  remarked  a s Eutamlas.  species  first  h a b i t a t use p a t t e r n s .  Chapter  known  i t s geographically  social  the p o p u l a t i o n .  animals'  (1972)  investigated  and s o c i a l  minimus  on a l t i t u d i n a l  presents and  formerly  chipmunk  chipmunk a n d  preference  in  with  have  1971; Meredith,  These  preference  least  use  a t Kluane Lake i n  and M e r e d i t h  studies  Tamias  1 9 7 1 ; Sheppard,  Chappell,  (Tamias minimus)  preference  group of western chipmunks these  space  a v a i l a b l e on w e s t e r n s p e c i e s  t i m e , a number o f  space  affecting  (1969),  Sheppard  paucity of data  that  INTRODUCTION  Yukon.  (1966),  Forbes  GENERAL  investigates factors least  southwest  1:  then  Population  compared  examines the  t h e h a b i t a t and i n v e s t i g a t e s t h e  to  those  levels,  home r a n g e and  reported  i n previous  r e l a t i o n s h i p between s o c i a l  spacing  2 and  space  whether dominance  use.  To d e t e r m i n e  chipmunks is  Chapter resulting availability then  defend  affected  chipmunks a v o i d 4 from of  whether  by  encountering evaluates  these  exclusive ownership each  whether  trapping  and  resources  compare t r a p a n d t e l e m e t r y  range.  social  spacing  areas, of  occurs,  whether  space,  and  I ask social  whether  other. t h e p r o v i s i o n of food pre-baiting enough t o a f f e c t  b a s e d methods o f  and c o v e r  increases  the  home r a n g e .  I  e s t i m a t i n g home  3 CHAPTER 2:  DEMOGRAPHY,  HABITAT USE, AND HOME RANGE OF THE LEAST  CHIPMUNK  IN THE SOUTHWEST YUKON  INTRODUCTION  This 1985  chapter  of  least  the southwest  habitat  with  those  since  1971;  provides The  and  1976;  rather  populations  Snyder,  and  than  common such  congeners  States,  describe  1974).  data  are  used  the  Lake  to assess  home  ranges,  a r e compared  range and h a b i t a t a s s o c i a t i o n s  used  1976;  in  the l i t e r a t u r e ,  estimates (Heller,  in assessing  197i;  Chappell,  Sheppard,  1978).  This  f o r comparison.  few s t u d i e s o f t h e l e a s t  Skryja,  chipmunk  have  opportunity  characteristics 1969;  o f home  i n t e r a c t i o n s with  competition  of a  minimus) near Kluane  habitat preferences  are relatively  studies  ample  t h e summers o f  reported.  chipmunk  Meredith,  during  r a n g e movements  chipmunks,  and d e r i v e d  estimates  several  competitive  of  previously  the l e a s t  (Tamias  The home r a n g e  preferences  Published  on  collected a n d home  chipmunks  Yukon.  demographic p a t t e r n s  of  data  a n d 1986 on t h e demography  population in  presents  chipmunk  morphological demography  t h a t have n o t f o c u s s e d traits  (Criddle,  and l i f e - h i s t o r y 1943; S h e p p a r d ,  However, s t u d i e s o f t h e demography o f o t h e r a r e more  1973; S u l l i v a n  numerous  e t al, 1 9 8 3 ) .  (Gashwiler, The chipmunk  1970; T r y o n population  4 near  my s t u d y a r e a  demographic 1984; too  studies  Gilbert,  has been  of other  patterns,  densities  reported  in  t o permit ln  previous  found  near  monitored  s m a l l mammals  unpublished data).  short a period  previously  a t Kluane  Although  since  (Gilbert my f i e l d  study  studies  of  can  be  and  covered  demographic  compared  chipmunks,  ln  and K r e b s ,  work  r o b u s t c o n c l u s i o n s about  this  1980  t o those  with  those  my s t u d y a r e a a t K l u a n e .  METHODS  Study  area  The 138°  s t u d y a r e a was l o c a t e d  W).  3 km  Two t r a p p i n g g r i d s  from t h e  Creek  on  the  l a k e on  forest  (Picea  were e s t a b l i s h e d  and  glauca),  open s p r u c e  to  open  bounded  bounded  ranged  to  Lake  from  2 and  by S i l v e r  by t h e A l a s k a closed  - soapberry  meadows  ( 6 1 ° N,  approximately  fan,  partially  Vegetation cover  shrub-land,  canadensis)  e a s t of Kluane  an o l d a l l u v i a l  north-east  Highway on t h e s o u t h .  just  dominated  spruce  {Shepherdla by Dryas  d r ummondi 1.  Live-trapping  Population trapping.  size  a n d home  ranges  were  monitored  using  live  I n 1985 I s e t up two r e c t a n g u l a r t r a p p i n g g r i d s ,  6 ha  5 in  size  traps of  at  day  apart.  During  of  seeds,  3  for  total  of  marked  with  in  capture  location,  by D  weight,  age  of  according  and  classified of  the I  whether  each  they  to mid-September,  (Figure  in order  to  increased  2.1).  nest In  sites  May,  the  distance  third  of  grids grid  (adult  (Chapter  or  I  most  size  day of  3. day  pelage I  was  recorded  juvenile)  and  breeding  (small,  medium or  pregnant. to  of  Males the  were  position  respectively. to  X)  late  August,  In  mid-July,  f r o m 6 ha t o first  local  4).  to I  it  removed  westward.  km  to traps  Home  and  1985,  to  I  that  entered  home r a n g e s . 1  1985,  8 ha so  expanded both g r i d s  grids  X and e x p a n d e d  the  chipmunks  assessed  when j u v e n i l e s  1986,  between  of  their  I  pre-  during  Captured  and  1986.  (grid  c o m p l e t e l y encompass  movement between eastern  most  left  i n the morning  according  e a r l y May  grid  traps  identification.  non-breeding  from  mixture  every 2 to  morning  and  obviously  or a b d o m i n a l )  would c o n t a i n  traps  were  (scrotal  May  2,  nipple  testes  expanded the western-most it  to  or  from e a r l y  day  chipmunk.  breeding  grids  set  on t h e  class  as  trapped  I  per s e s s i o n .  sex,  condition  females  of  visual  of  with a  t h e a f t e r n o o n and e v e n i n g  for  condition  Longworth  l o c k e d open a n d  toe-clipping  breeding  in  t r a p p e d each g r i d  them open  6 checks  Nyanzol  were b a i t e d  sessions,  checked  were p e r m a n e n t l y marked  large)  I  a f t e r n o o n and e v e n i n g a  were c a p t u r e d  and were  sessions.  locked  generally  morning,  Traps  trapping  d a y 1 and  were  Animals  21 m a p a r t .  between t r a p p i n g  morning  1,  m  and s u n f l o w e r  weeks.  Traps  500  stations  oats  balted 3  and  the  18 ha I  also  eliminate from the range  6  Fig. and text  2.1 X. for  —  The  Shading  study area, indicates  details).  showing  extent  of  location grids  at  of  trapping  grids  d i f f e r e n t times  H  (see  7  estimates of  from  sampling, In  (see  1986 t h e two g r i d s  September,  September.  Table YEAR  4 ) . Both  The  2.1—Summary  21 m  1986  42 m  X  alone  1 km  level  H)  alone  was  trapped  from  from m i d - J u l y t o  trap-spacing,  a r e summarized  the season  l n e a r l y May and m i d -  was t r a p p e d  locations,  GRID  .5 km  trapped  (grid  of g r i d - t r a p p i n g  SPACING TRAPS GRIDS  1985  were  grid  grid  o f t h e two g r i d s  a coarser  was i n c r e a s e d t o 42 m.  grids  sizes,  enough t o w a r r a n t  were t r a p p e d f o r o n l y h a l f  the easternmost  m i d - J u l y , and  periods  high  so t r a p - s p a c i n g  Chapter  May t o  1985 were  and t r a p p i n g  in Table 2.1.  changes  SIZE  PERIOD OF OPERATION  H  6 ha  H  May t h r u A u g .  X  6 ha t o J u l y 17, t h e n 8 ha  X  May t h r u A u g .  H  18 ha  H  May t h r u J u l y 16 and m i d - S e p t .  X  18 ha  X  e a r l y May a n d J u l y 16 t h r u Sept.  Radio-telemetry  R a d i o - t e l e m e t r y was residents continuous  i n both y e a r s . estimates  collar-mounted;  of  used t o l o c a t e In  1986 I  home  each package  used  range.  weighed  nest-sites  o f most  telemetry  to provide  Radio-transmitters  2 - 2.5 g ,  less  adult  than  were 5% o f  9 mean 1986 to  a d u l t body w e i g h t  permit  triangulation  were c o l l e c t e d route  on  source  and  accustomed long  as  t o my did  undisturbed over  a  1 minute  eating. if  not  animal,  person  and  was  Habitat  was  doing  to evaluate  5  m.  of  readily  Upon  Its grew  animals  animal  tracking,  or  as  locating  substrate,  those  i f the  the  to  apparently undisturbed  the a s s o c i a t i o n  by  quadrat  m grid  substrate.  Open and  each other  species. estimates 10  quadrat. points,  type  data  was  an  behaviour which  were  disturbed,  i f more t h a n  one  classification  c h i p m u n k s and  radius  Chipmunks  location,  recorded  movements o f  from  food  range  t o t r a c k each s i g n a l  within the  home  low  I walked a r e g u l a r census  remained  I recorded  so  too  present.  In o r d e r  30  route  approach  p e r i o d and  assistant  locations,  location.  L o c a t i o n o n l y was  an  the  its  presence  S i g n a l s t r e n g t h was  sampling.  leaving  record  I  of animal  using scan  each g r i d ,  (51 g ) .  two  according  cover  the  recorded  differed was  mean p e r c e n t  cover  grid  stake  reassessed cover  less  as  more  30  type  m  and  were d i s t i n g u i s h e d  of  taken  by  each  dominant cover  communities  observers,  from  Where e s t i m a t e s  Quadrats with  was  different  m measured  percent  to the  closed forest  cover  home r a n g e  habitat, I classified  a c c o r d i n g to percent  Percent by  their  between the  the dominant t h e mean o f over  a  visual  circle  a t the c e n t r e  of  of  each  20  percentage  by d i f f e r e n t  observers.  than  than  tree  25%  were c l a s s e d as  10 open f o r e s t ;  those  with cover  of over  25%, and In which t r e e  canopies overlapped, were c l a s s e d as c l o s e d  forest.  I r e c o g n i z e d e i g h t d i f f e r e n t h a b i t a t types  Table  (Table 2.2).  2.2--Habitat c l a s s i f i c a t i o n s SUBSTRATE  HABITAT TYPE  SPECIES  1. Closed spruce -soapberry  Picea glauca,  2. C l o s e d spruce -moss  canadensis,  Arctostaphylos canadensis, Azctostaphylos  Fine  till  boreale,  uva-ursl  Plcea glauca, Lupinus  Shepherdia  Hedysarum  Shepherdia  Hedysarum rubra,  arcticus  boreale, A. u v a - u r s i ,  Soli, moss  3. Ecotone between open and c l o s e d habitats  Shepherdia canadensis, Eleagnus commutata,  4. Open p o p l a r , soapberry, spruce  Populus balsamlfera, Shepherdia canadensis, Picea glauca, Dryas drummondii, Arctostaphylos rubra, A. uva-ursi, Oxytropis campestris  Coarser till  5. Open dryas  Dryas  Coarser till  6. Open spruce -soapberry  Picea glauca, Shepherd la canadensis, Dryas drummondi1, Arctostaphylos rubra, A.  u v a - u r s l , Hedysarum  spruce  8. Coarse t i l l , soapberry  Picea  Shepherdia  glauca  Fine  till  boreale  drummondi1  uva-ursl, Oxytropis  7. Dead  Salix spp. Arctostaphylos  Coarser till  campestrIs  Very coarse till canadensis  Very coarse till  11 Five and  of these Wingate  spruce  conformed (1976).  (Picea  canadensis); balsam  two  open  flats.  layer  licorice  (Arctostaphylos shrub,  and  and  forest  communities,  almost  complete  soapberry  community  has a  soapberry  shrub,  open s p r u c e - s o a p b e r r y layer  dominated  campestris),  by  that  A.  bearberry, is  alluvial  dead  (drowned) s p r u c e ,  glauca),  communities  has a s p a r s e  similar  with  to  some s p a r s e  soapberry  The  with  Three  sparse  soapberry,  characterised berry  herb  additional  and an ecotone  and sometimes s i l v e r  and  the  closed  a denser little  moss.  bare  habitat  (Oxytropis o f t h e open types  alluvial  soapberry,  (Bleagnus  The  and a herb  locoweed layer  sprucelayer of  till  between t h e c l o s e d  by  herb  bearberry  layer,  Dryas,  Dryas  and  cover.  stony substrate  and  has a s u b s t r a t e o f c o a r s e r t i l l ,  bare  forest  s p r u c e - s o a p b e r r y and  boreale)  sparser herb  (Shepherdla  soapberry;  uva-ursi),  moss  p o p l a r - s o a p b e r r y community. till  -  of Krebs  communities,  spruce-soapberry  (Hedysarum  and  and a  forest  s p r u c e - m o s s community  root  rubra  two c l o s e d  balsamifera)  The  classifications  -moss  (Populus  drummondii  habitat  T h e s e were:  glauca)  poplar  of  to the  with  a n d open  willow  commutata).  were:  (Salix  12 RESULTS  Demography  Jolly-Seber sizes  from  the  e s t i m a t i o n was  used  capture-recapture  data  results  are p l o t t e d  animals  did  not d i f f e r  ( t = 2 . 0 0 , p=.10). revealed  no  grid  of a d u l t s  summer  than  i n 1985  16.2  i n 1986).  of  to  fact  grid  than  of  grid  i n 1986  both  after  May,  since  grids  X  t h e r e was  although  on  both  grids  occupying  year  numbers The  expanded  of g r i d  H  of g r i d  t r a p p i n g stopped  H  the  the  spring  number end  i n 1986 (17.2  of  of the  (23.4)  in  1985,  the  size  of a d u l t s  should  asymmetry  l n the  grids  c o u l d be  due  I n c l u d e d more f a v o u r a b l e X.  p o p u l a t i o n peak e v i d e n t No  The  increase*ln  i n 1986. the  both y e a r s .  w i t h the  a three-fold  to the next,  22.6  i n 1985  ln either  t-test  of  through  grid  X  animals  average  over  coincident  year  emergence.  an  of  grid  a  number o f  stable  were more a d u l t s  the expansion  juvenile  was  H and  grids,  the  These  mean number  grid  both  ln  remained  the expansion  mid-summer  result H  animals  that  types  The  one  h i g h e r on  o f new  habitat  Since  from  number the  on  (13.7) o r on g r i d  the g r i d s  have been  There  from  1982).  the  between  There  both g r i d s ,  slightly  season.  p=.07).  each on  data  (Seber,  1985,  difference  present  on  season  males dropped mating  pooled  (t=1.91,  chipmunks per  and  On  In  significantly  significant  between y e a r s  number  i n F i g u r e 2.2.  to calculate population  l n F i g u r e 2.2  p o p u l a t i o n peak before  i s the  i s shown f o r  juveniles  were o l d  13  Fig.  2.2 -- J o l l y - S e b e r p o p u l a t i o n e s t i m a t e s  s  f o r G r i d s X and  H  14  May 20  June July 1985  Aug  May June July Aug 1986  Sept  Grid H O—O  15+  females  O  m a l e s  //°  \ 5+  J  .  May  —  1  June July 1985  1  1  Aug  \  \  1  May  1  June 1986  h  July  15 enough t o enter t r a p s .  The appearance of  consistent  to  from  year  minimus, breed o n l y 1966;  once a year  Sheppard, 1 9 6 9 ) .  A p r i l - e a r l y May, '• b i r t h emergence  in July.  indicated  Table GRID  X  h  i n Canada  in  late  including  ( C r i d d l e , 1943;  That  May-early  there  Tamias Forbes,  June,  of j u v e n i l e s  captured  have a range of a p p r o x i m a t e l y 10 g.  but t h i s d i f f e r e n c e  and j u v e n i l e  i s some v a r i a t i o n i n t h i s i s  emerged on g r i d  i n the  From Table 2.3  X than  on g r i d H  i s not s i g n i f i c a n t ( t = 1 . 0 3 ,  p>.50).  2 . 3 - - P o p u l a t l o n means YEAR  NUMBER OF ADULTS  NUMBER OF JUVENILES  FEMALES  FEMALES  HALES  MALES  19 8 5  MEAN = S.D. =  6-2 2.0  11.1 1.8  1986  MEAN = S-. D . =  7. 8 2.7  8. 4 1.5  •9 . 3 3.2  3.6 1.7  19 8 5  MEAN = S.D. =  5. 5 0. 8  8.2 1.2  5.8 1. 7  3. 7 1. 0  19 8 6  MEAN = S.D. =  12.1 0. 8  11. 4 0.9  1.7* 0 *  0* 0*  •sample size too low to a c c u r a t e l y d i s c o n t i n u e d a f t e r j u v e n i l e emergence). Among a d u l t s , males  i n J u l y was  Mating tends t o occur a l l a t once i n l a t e  appears t h a t more j u v e n i l e s  in 1985,  Chipmunks,  by the f a c t t h a t weights  same s e s s i o n it  year.  juveniles  the sex  i n 1985  calculate  4.1 1.6  N  (trapping  r a t i o vas c o n s i s t e n t l y biase'd  (1.57 males/female) on g r i d X.  toward males  ' 7.8 1.1  On g r i d  toward  H i t was biased  (1.50 males/female) but wa3 n e a r l y  even i n  16 1986  (0.95 m a l e s / f e m a l e ) .  Among  m a l e s were c a p t u r e d on b o t h Mean are  minimum  presented  consistently meaningful  s u r v i v a l rates Table  exceeded  (Boonstra,  GRID  X  h  X  h  Adult 0.89 .  2.4. unity  between-sesslons Jolly-Seber  generally  females  (three-weeks)  not b i o l o g i c a l l y  1985).  Rates  YEAR  AGE  SEX  1985 1986  adult adult  male male  0.88 0.82  1985 1986  adult adult  female female  0.94 0.86  1985 1986  adult adult  male male  0.89 0.93  1985 1986  adult adult  female female  0.95 0.86  1985 1986  juvenile juvenile  male male  0.34 0.38  1985 1986  juvenlle juvenile  female female  0.60 0.87  1985 1985  juvenile juvenile  male female  0.67 1.00  survival  the sexes. lower  than  survival estimates  and a r e t h e r e f o r e  remained  high  over  the  T h e r e were no d i f f e r e n c e s i n a d u l t  between  more  grids.  2 . 4 - -Mean Minimum S u r v i v a l  Table  or  in  juveniles,  Juvenile  f o r males  SURVIVAL  summer, a v e r a g i n g  s u r v i v a l between  s u r v i v a l averaged  (0.44) t h a n  f o r females  grids  0.6, b u t was (0.84).  For  17 1985,  the  lower  on  year grid  were c a p t u r e d consistent breeding on  grid  with X  t h e most  than  on g r i d  with  females  on g r i d X  in  data, H.  Nearly  1985  the d i f f e r e n t  juvenile  as  number  twice  on  a s many  juveniles  grid  H  (22  v s .13),  females.  X, w h i l e  Seven  4 were r e s i d e n t  H i n 1985.  survived  b e t t e r than  juveniles.  On  while  on g r i d  X  Table  2.5—Minimum  survival  is listed  m a l e s on b o t h  grid  grids,  adult adult juvenile juvenile  male female male female  X X  adult adult juvenile juvenile  male female male female  X X  Adult weight and  averaged did  50.4 g o v e r  not  differ  W e i g h t s were l o w when a n i m a l s and I n c r e a s e d  g on a v e r a g e  juveniles,  female while  weight  lactating  SURVIVAL  both  were f i r s t  remained increased. than  0.75 1.00 0.60 1.00  .  grids  trapped  of over  between i n May  stable  (8) (5) (5) (7)  years years.  (mean =  50 g by e a r l y  relatively  (N)  0.56 ( 9 ) 0.75 ( 4 ) 0.67 ( 3 ) 1.00 ( 6 )  and b o t h  significantly  t o an average  p o i n t , male w e i g h t  summer, w h i l e  f o r a d u l t s and f o r  over-winter s u r v i v a l  h h h h  From t h i s  Females  t h e r e v e r s e was t r u e .  SEX  g),  2.5.  H, a d u l t s s u r v i v e d b e t t e r t h a n  AGE  2.6)  i n Table both  GRID  44.9  was much  of breeding  were r e s i d e n t on g r i d  Minimum o v e r w i n t e r  (Table  survival  June.  over the  F e m a l e s were h e a v i e r b y 5  d u r i n g pregnancy.  The s m a l l e s t  18 juveniles weight. differ  to  enter  By  t r a p s weighed  September,  f r o m mean a d u l t  Weights  used  mean  juvenile  weight  (50.4 g ) .  to c a l c u l a t e  by w e i g h i n g e a c h c a p t u r e d are  up  t o seven  2.6—Means  means  weight  from e n t i r e l y  grew t o a d u l t  (49.7 g)  d i d not  i n T a b l e 2.6 were o b t a i n e d  a n i m a l once  repeated weighings  means a r e n o t d e r i v e d  Table  29 g, b u t r a p i d l y  per t r a p - s e s s i o n .  There  per animal per year, so the independent  measurements.  ( a n d r a n g e s ) o f chipmunk w e i g h t s  ( i n g)  YEAR  ADULT MALES  ADULT FEMALES  JUVENILES  1985  47.2 (37 - 57)  53.1 (40 - 65)  44.3 (31 - 52)  1986  47.8 (39 - 57)  52.6 (39 - 64)  48.0 (29 - 53)  When breeding 1986,  live-trapping condition  breeding  was u n d e r w a y  c h a s e s a t t h e end  buried  In snow.  and  in  e a r l y June of August.  This  30-day g e s t a t i o n  1985,  each g r i d  1986,  t h e same r e s i d e n t  not breed.  April  males the  when  a l l males  They  for  had one r e s i d e n t  of  May.  I observed  the  traps  In  several  were  still  c a p t u r e d i n May were i n in late  had f i n i s h e d  agrees with C r l d d l e ' s  period  were a l r e a d y i n  end  April.  began l a c t a t i n g  In 1986.  of a  did  by l a t e  of  Females  May,  so u n t i l  In both y e a r s ,  condition.  beginning  in  and remained  mating  breeding  began  May  lactating  i n 1985 by t h e  (1943) e s t i m a t e  Tamlas minimus bozealis.  female  f e m a l e on g r i d  that  d i d not  H was t h e o n l y  For both sexes, a l l of the  yearlings  breed. female  In In that  c a p t u r e d on  19 the  grids  contrasts  bred with in  following  their  western  Juveniles  10  estimate  were  panamintinus presented  (1969)  first  breeding found  d i d not  observed  season.  breed  This  Tamlas minimus  that  in  the  spring  This  juvenile  supports  here  o f young  at  No the  juveniles  entered  Hirshfeld  emergence  because  on J u n e 2 9 ,  on t h e s t u d y a r e a  Juveniles f i r s t  and T. p a l m e r ! .  trappability  who  Alberta  30, 1986.  years. of  first  birth.  a n d on J u n e  i n both  their  Sheppard  oreocetes  1985  during  5  t r a p s on J u l y (1977)  and B r a d l e y ' s  Tamlas  weeks o f a g e f o r  estimates  of  recruitment are  expansion  of  grids  precluded  that  and  low  analysis.  Home Range  Home r a n g e was e s t i m a t e d animals  ln  collars,  calculations.  Female  developed  108  a severe  infection  other (and  to  telemetry  grid  H)  data  were a c r o s s from  that  Silver  with  because Creek  a l l analysis  might  for  have  from t h e  because she  been c a u s e d  were t o o t i g h t  f o r which t h e d a t a  Except and  were a l s o  when I  from  nearly  on h e r neck due t o t h e t i g h t n e s s o f h e r c o l l a r . t h e end o f t h e summer,  23  radio-  were e x c l u d e d  60 was e x c l u d e d  excluded  remove t h e c o l l a r .  collars  from  Female  was  the  c h i p m u n k s were f i t t e d  eye i n f e c t i o n  became t r a p - s h y u n t i l able  females  of her t e l e m e t r y l o c a t i o n s  grid.  an  Twenty-five  b u t two ( b o t h  home r a n g e half  1986.  from  was  by She  finally  i n e a r l y May when some o f t h e  caused  chafing  excluded),  around  the other  t h e neck chipmunks  20 did  not appear  to  suffer  of  1238  adverse  effects  from  wearing  radio-  collars. A  total  through sizes  telemetry  S e p t e m b e r , were  were c a l c u l a t e d  on a minimum o f 20 home r a n g e .  used  chipmunk.  w i t h a mean a r e a  et a l  w i t h an  other  home r a n g e s .  average  (1980) f o u n d  that  Home r a n g e o f 4.86  ha  areas  range  (S.D.=2.35).  o f chipmunk  home r a n g e  May  Home  range  method,  based  54 p o i n t s p e r  home r a n g e 0.97  Home  significantly  from  20 c a p t u r e s was  from  (mean=2.89 h a , S.D.=1.331), t=4.34,  aspects  of  to a c c u r a t e l y estimate  (mean=6.13 h a , S.D.=1.96) were females  collected  b y t h e minimum c o n v e x p o l y g o n  minimum number n e c e s s a r y eastern  to estimate  locations,  Mares  locations,  of the  ha t o 9.65  ranges  larger  ha  o f males  than  p<.001).  i n appear  the  those  of  A n a l y s i s of  i n Chapters  3 and  4.  Habitat  To  preference  assess  classification 2.3).  data  H  preference,  are l i s t e d  i n Table  X, a l l 8 h a b i t a t  only 4  open p o p l a r - s o a p b e r r y the  for  associations  the  live-trapping  closed  and Dryas  telemetry location  map  I used  f o r each  habitat grid (Pig  spruce-soapberry,  flats.  type  the  represented, while  Both  d a t a were p l o t t e d  between h a b i t a t data,  used  2.2.  types are  o f them o c c u r :  and  I  to construct a habitat  Habitat types On g r i d  grid  habitat  and  a G test  the  t o compare  ecotone,  live-trapping  on t h i s  frequency  on  of the  map  to  test  use.  For  frequency  21  Fig.  2.3  H a b i t a t maps: 1. C l o s e d  (a)Grid  spruce  2. C l o s e d s p r u c e 3.  soapberry moss  Ecotone  4 . Open p o p l a r 5. D r y a s  7. Dead  soapberry  flats  6. Open s p r u c e  soapberry  spruce  8. Open t i l l  X,  soapberry  (b)Grid  H  22  23 of  capture  the  In e a c h  v e g e t a t i v e community w i t h t h a t  p r o p o r t i o n of  performed  separately  grid,  since  after  each  grid  p<.001 i n  H,  both  frequently  years.  in  in  on  Dryas  t o changes For  habitat  In  traps  and  but  in  the  size  habitat  significantly  of  each  differed  In t h e X,  G  type  with  sampling tests  .001<p<.005.  For  soapberry habitat When c l o s e d captures  eight  was  and  captured and  more  expected  from  type  less  equal  often  than  than expected  in  not r e p r e s e n t e d i n  y e a r s were t h e r e f o r e  frequency of capture of  sizes. and  For dead  .025<p<.05.  habitat 1986  types grid  traps  from  t h e r e were grid  spruce  habitats  were  the expanded  1985  For  occurred,  habitat  there than  set  i n each  t h e s m a l l e r 1985  G=22.107, df=7  which c l o s e d  G=198.2, df=6 and  spruce-shepherdia were l o c a t e d  than  b u t more o f t e n  number  grid  absent,  habitat  were c a p t u r e d  comparing  the  0=22.945,  distribution.  relative  df=5  f o r 1986  trap  f r e q u e n c i e s were  Changes between  a l l three  G=12.107,  habitat  the o n l y h a b i t a t  H.  from  c h i p m u n k s were  expected  chipmunks  was  of g r i d  on  and  meadow i n 1985,  on w h i c h a l l  fewer  f o r each  spruce-shepherdia  from which the p o p l a r - s o a p b e r r y  grid  T e s t s were  and  both y e a r s ,  Observed  grid  significant  absent,  grids  open p o p l a r - s h e p h e r d i a  ecotone,  the expansion due  community.  differed  closed  Dzyas meadow  1986.  of  use  trap d i s t r i b u t i o n .  expected  that  i n b o t h y e a r s : G=23.578 f o r 1985;  frequently  the  in  f o r t h e two  the d i s t r i b u t i o n  distribution  for  set  from  expansion.  For  less  traps  expected  spruce-  p<.001.  occurred  expected.  For  on  grid  X  the  closed  spruce-moss h a b i t a t , to  the  was  expected  captures  the g r i d ,  in  Dryas  captures  than  t h e number  i n ecotone  medium v e r s i o n s large  version  greater spruce more  than  summarized observed  i n Table  number  of  ecotone  than  grid,  but The  number  for 2.7.  A + H  i s less  open  habitat  (habitats and d e a d  types  1 and 2 ) . spruce  the g r i d  habitat,  and f a r  expected.  the  f o r the grid.  i n the  habitat  had a  medium  number  a "-" i n d i c a t e s  expected,  the t i l l - d e a d  These  that  s m a l l and  expected  till-soapberry  expected  "0" means  grid  and  results are of captures  that  observed  that  observed  number, and t h e a b s e n c e o f  Trapping r e s u l t s  show t h a t  (4,5,6,  and  8) o v e r  Results  are  Inconclusive  habitats  sizes  The number o f  in  g r e a t e r than  indicates  than  by a " . " .  After  For a l l three  expected  large  equal  open s p r u c e - s o a p b e r r y and  expected  M  X.  of captures, while  the  captures equalled  i s indicated  forest  less  in  than  g r e a t e r than expected,  of captures  prefer  was  c a p t u r e s than  number  data  greater  expected  is  occurred there.  was  expected  grid  spruce-moss  of captures  of the  o f c a p t u r e s was n e a r l y  smallest ln  of the g r i d .  had f e w e r than  was  expected  habitat  number  on t h e  e x p a n d e d , more o f i t  fewer of  number  observed  (3 and 7 ) .  chipmunks  closed  spruce f o r the  I  25  2.7—Summary  Table  of  habitat  preferences  from t r a p p i n g  data.  HABITAT TYPE 1  H H X X X  85 86 85, 6ha 85, 8ha 86  The  tests  grids.  •  0 0  + +  +  +  + +  + +  similar  to  the  (type  3),  strong  while  positive  soapberry  (types  observed  and  trapping  locations  5) do  with  •  •  *  •  •  + 0 +  + +  locations  non-randomly  telemetry The  with  i n the  Dryas  telemetry  from  the  but  (type  with  and  for  and 2).  ecotone  There  open  number  were  association  inconclusive. flats  are  Grid  (types 6  associated  both  results  trapping results.  trapping data,  not d i f f e r  on  results  c l o s e d spruce-moss  were  inside  locations  show a p o s i t i v e  positively  results  association 4 and  both  8  numbers o f  the  the  7  open t i l l - s o a p b e r r y  association were  used  f o r 1986.  c o n s i s t e n t with  telemetry results  a negative  H  6  telemetry  expected  grid  trapping results  However, t e l e m e t r y d a t a  data  For  on  were  and  open s p r u c e - s o a p b e r r y  8) and  --  were p e r f o r m e d  2.4.  X were l e s s  X t r a p p i n g and  :  Observed  in Figure  with  5  Habitat types  (p<.001).  grid  4  -  •  plotted  for  > 3  0  same  trapping grids  -  2  is a  poplar-  telemetry  expected.  26  Fig. vs.  2.4  —  Observed  and e x p e c t e d  number  vegetation type. 1. C l o s e d 2. 3.  spruce  Closed spruce  soapberry moss  Ecotone  4.  Open p o p l a r  5.  Dryas  6.  Open s p r u c e  7.  Dead  8.  Open t i l l  soapberry  flats soapberry  spruce soapberry  of  telemetry  locations  27  Grid H (ZD observed E53 expected  Habitat Type  Grid X I I observed ESS expected  2  3  nfl 4  nl  5  Habitat Type  6  DM. 7  8  9  28 I compared  the h a b i t a t  p r e f e r e n c e s of a d u l t s and  u s i n g the r e c a p t u r e d a t a f o r 1985, were  most  numerous  (Figure  c a l c u l a t e d by d i v i d i n g the given  habitat  habitat.  by  the  Juveniles  h a b i t a t types  2.5).  such as  The  percent  percent  had  when j u v e n i l e  a  of  of  moss and c l o s e d s p r u c e - s o a p b e r r y  captures  index  observed  expected  proportion  dead spruce,  r e c a p t u r e data  h a b i t a t use  captures  greater  juveniles  of  soapberry and  was in a  i n that  captures till,  In  spruce-  than a d u l t s .  DISCUSSION  Demography  I found no evidence in  numbers  (1950) who of  of  least  of pronounced chipmunks  chipmunks.  (1973) who  reported  chipmunks.  Their  This  a  3-4  data  not support  evidence  Two  only  (Gashwiler,1970;  typical  with Tryon" and  population  s i n c e peak p o p u l a t i o n s were o n l y 2-3 populations.  f l u c t u a t i o n s are  contrasts  year  did  fluctuation  a t Kluane c o n t r a r y to C a l l a h a n  a s s e r t s t h a t wide p o p u l a t i o n  western  year-to-year  cycle  in eastern  t h e i r c l a i m , however,  times the s i z e of the  demographic s t u d i e s of Tamias townsendl1 of  2-  to  3-fold  S u l l i v a n et a l , 1983).  Snyder  annual  lowest found  fluctuations  29  Fig.  2.5  habitat observed  —  type.  Habitat Value  index  of index  / % captures 1. C l o s e d  use  f o r each  expected.  spruce  2. C l o s e d s p r u c e  soapberry moss  3. E c o t o n e 4. Open p o p l a r 5. D r y a s  7. Dead  soapberry  flats  6. Open s p r u c e  soapberry  spruce  8. Open t i l l  for  soapberry  adults habitat  and j u v e n i l e s v s . type  = %captures  Habitat Use Index  Habitat Use Index  31  present  study  i tis  impossible to t e l l  ecotone,  avoid ecotone  reflects  a  than  problem  indecision  more c o n s i s t e n t according  Home  or a r e I n d i f f e r e n t  w i t h my h a b i t a t  on t h e p a r t  120  (Table  home r a n g e times  square  of least  scheme,  R e s u l t s may further  rather  have  been  Into c a t e g o r i e s  bordered i t .  study.  chipmunk  Different  methods  since  the  t h e minimum In  sufficiently  minimum  most  of  clumped  here,  these  that  so t h i s  and g r i d  home r a n g e  method used  studies,  of  capture  the estimated  were  since  t h e methods  Feldhamer low  these squares  estimates  points  were  method  would  area t o  used  areas  In t h e p r e s e n t  does n o t p r e c l u d e c o m p a r i s o n .  on sample s i z e .  w h i c h sample s i z e s  complicate  method  smaller  were 4  i n other  use o f t h e c o n v e x p o l y g o n  p r e s e n t s more o f a p r o b l e m , dependent  used  area  yield  i n t h e Yukon  home r a n g e s  convex polygon  have added o n l y a f r a c t i o n listed  chipmunks  of l e a s t  a r e a method g e n e r a l l y  t h a n does  the figure Sample  here  size  are a l l  (1979) was t h e o n l y s t u d y i n  enough  to  severely underestimalte  range. In most  adults range  ecotone  types that  sizes  size  2.8).  comparisons,  home  This probably  classification  o f chipmunks.  l f I had d i v i d e d  to the habitat  to i t .  select  range The  to  whether chipmunks  c a s e s , home  range  areas  o n l y , as i n the present study. areas  manipulation,  were r e p o r t e d home r a n g e s  during the listed  listed  are  For studies course  t h e means f o r i n w h i c h home  o f an e x p e r i m e n t a l  here a r e f o r t h e c o n t r o l  area.  32  Table  2.8—Chipmunk  home r a n g e s  from t h i s  and p r e v i o u s s t u d i e s  STUDY, LOCATION  SPECIES  METHOD  B l a i r 1942 Michigan  Tamias s t r i a t u s  Trapping. Minimum c o n v e x p o l y g o n (MCP).  0. 90  Broadbrooks 1970 Washington  T.  amoenus  Trapping. No. t r a p s used * trap-unit area.  1.,20  B u r t 1940 Michigan  T.  striatus  Trapping. MCP.  0. 62  C h a p p e l l 1978 California  T. T. T.  minimus amoenus speciosus  Trapping. No. t r a p s used * trap-unit area.  1. 05 1. 31 1. 39  E l l i o t t 1978 New York  T. s t r i a t u s  O b s e r v a t i o n . No. 0. 17 quad, u s e d * a r e a .  F e l d h a m e r 1979 Oregon  T. minimus  Trapping. 0. 04 Standard diameter.  F o r s y t h and Smith 1973,  T. s t r i a t u s  Trapping. Minimum a r e a .  0. 18  G l e n n i e , 1988 Yukon  T. minimus  Telemetry. MCP.  4. 86  Mares e t a l 1976, Penn.  T. s t r i a t u s  Trapping. MCP.  0. 11  Mares e t a l 1982, Penn.  T. s t r i a t u s  Trapping. Minimum a r e a .  0. 09  M a r t i n s e n 1968 Montana  T. minimus T. amoenus  Trapping. MCP.  1. 14 0. 86  S h e p p a r d 1972 Alberta  T. minimus T. amoenus  Trapping. Minimum a r e a .  0. 85 0. 99  S t a t e s 1976 Oregon  T. amoenus  Trapping. Boundary s t r i p .  0. 75  V e r g e r 1953 New York  T. striatus  O b s e r v a t i o n and Trapping. MCP.  0. 07  Ont.  HOME RANGE (ha)  33  T h e r e have been parameters  on  Eisenberg, was  (where and  that  (1983)  omnivores.  mass  is  of  early  derived  i n grams).  i s 4.86 h a ,  for  a  50 g  by  Mace  12 t i m e s  and  Harvey's Table  allometric  principle  ranges.  Of  2.6 r e v e a l s that  a l l the  larger  also  heaviest  from the a l l o m e t r i c Harvey  listed  home r a n g e s o n l y weigh  mass: A=2.70M0.63  mass  i n kilograms). equations  scales size  g r a n i v o r e s , and  as  0.007M1.12  of  Tamlas  (about  here.  granivores,  Western  than t h e i r about  Home  minimus a t  that  home r a n g e .  These  ha.  not f o l l o w the  listed,  larger the  home  eastern  home r a n g e , b u t i t  range  size  f r o m McNab,  listed  allometric  predictions 1.23 ha f r o m  t h e home  here  congeners, u s u a l l y the  predicted  0.56  have  g e n e r a l l y exceed  chipmunks  eastern  50 g, s o  underestimates of  100 g ) .  e q u a t i o n s , 0.63 ha  for  does  animals should  has t h e s m a l l e s t  (where  by McNab's e q u a t i o n  Tamias  species  f o r the  They s e p a r a t e d  for granivores, that  Mace  granivorous so t h e i r  predicted  chipmunk  Tamlas strlatus,  sizes  that  equation  chipmunk,  Mace and  t o body  a n i m a l , 0.409 ha, and i t i s 8.8 t i m e s t h a t  Examining  the  (1963) i n w h i c h he  largely  home r a n g e  work  McNab  herbivores,  ( i n ha)  The  Much o f t h i s  f o r rodents.  are  life-history  ( B l u e w e i s s e t a l , 1978;  allometric  groups:  size  predict  1979).  by  t o body s i z e  chipmunks  home r a n g e  Kluane  paper  i n hectares,  trophic  Yukon  to  body s i z e  was r e l a t e d  have  home r a n g e  three  predicted  an  attempts  1983; W e s t e r n ,  home r a n g e  of  into  basis  i s expressed  Harvey  taxa  by  area  relation  the  1981; P e t e r s ,  inspired  reported  several  have  range larger  over 1 ha, but  equations provide  discrepancies  indicate  that  is  34 factors  such  as  d e n s i t y outweigh  latitude, the  influence  McNab approach has l i t t l e be g e n e r a l l y detailed  food  availability,  of body s i z e .  predictive value,  true at the taxonomic l e v e l  information is  life-history  needed  to  and p o p u l a t i o n  For t h i s  and t h i s  is  genus the likely  to  of the genus where more  account  for  differences  in  characteristics.  SUMMARY  I compared population studies  of  of  demography, h a b i t a t least  chipmunks  chipmunks.  previous s t u d i e s of t h i s year-to-year  than  Population and  other  Kluane size  Lake is  species,  but  any r e p o r t e d for the genus. the  conducted  habitat  at  distant  Nevada, C a l i f o r n i a ;  found  H a b i t a t use comparison  by  different  the  Home range s i z e s are at l e a s t 4 times  limited  the mountains  with p r e v i o u s  lower than i n most  chipmunk  with previous s t u d i e s i s in  home range of a  s t a b i l i t y of the p o p u l a t i o n i s s i m i l a r to that  i n other chipmunk s p e c i e s . greater  at  choice and  the c e n t r a l  of Oregon).  I  s h r u b - l a n d and open f o r e s t  to  fact  that  they were  locations  Rocky Mountains  (Sierra  i n A l b e r t a ; and  found that Yukon chipmunks p r e f e r closed  g e n e r a l l y c o n s i s t e n t with the f i n d i n g s  canopy  forest,  of previous  which  studies.  is  35  There  have  been  several  parameters  on t h e b a s i s  Elsenberg,  1981; P e t e r s ,  was  inspired  reported (where and  by an  that  body  relation  early  paper  home r a n g e  by  (1983)  o f home r a n g e  derived  trophic  omnivores.  Yukon  chipmunks  predicted is  Kluane  home  range  i n grams).  is  size The  4.86 h a ,  are  mass i n  home r a n g e that  for  a 50 g a n i m a l , 0.409 h a , a n d I t i s  by  Mace  and  Examining  Harvey's  equation  T a b l e 2.6 r e v e a l s  allometric  principle  ranges.  Of  that  a l l the  species  has t h e s m a l l e s t  also  heaviest  100 g ) .  from the  allometric  Mace a n d H a r v e y  sizes  listed  equations,  f o r granlvores,  here.  home r a n g e s t h a n t h e i r o n l y weigh  about  underestimates  50  Home  that  g, s o t h a t  o f home r a n g e .  does  0.56  not  ha.  f o l l o w the larger  the  home  eastern  home r a n g e , b u t i t i s range  size  exceed  usually  the a l l o m e t r i c  These  at  predicted  predictions 1.23 ha f r o m  the  home  western chipmunks l i s t e d here have eastern congeners,  (where  1  b y McNab's e q u a t i o n  0.63 ha f r o m McNab, generally  so t h e i r  Tamias minimus  listed,  Tamias strlatus, (about  of  8.8 t i m e s  chipmunk, the  g r a n l v o r e s , and  a n i m a l s s h o u l d have  chipmunk  Mace  They s e p a r a t e d  1  Tamias  a  equations f o r the  f o r granlvores,  that  larger  A=2.70M°-*  as 0.007M - *  predicted  work  w h i c h he  granlvorous  size  1978;  kilograms).  herbivores,  ( i n ha) s c a l e s  12 t i m e s  in  t o body mass:  largely  al,  et  (1963)  f o r rodents.  groups:  life-history  Much o f t h i s  allometric  t o body s i z e  three  1979).  McNab  was r e l a t e d  have  to predict  (Blueweiss  1983; W e s t e r n ,  into  mass  size  area i s expressed i n hectares,  Harvey  taxa  of  attempts  over  range  larger  1 ha, but  equations provide  discrepancies  indicate  that  36 factors density  such  as  outweigh  latitude,  the Influence  McNab a p p r o a c h h a s l i t t l e be  generally  detailed  food  true  availability,  o f body s i z e .  predictive value,  a t t h e taxonomlc  information  is  level  needed  to  and p o p u l a t i o n  For this and t h i s  i s likely to  o f t h e genus  account  genus t h e  where more  for differences i n  life-history characteristics.  SUMMARY  I compared demography, h a b i t a t population studies  of  studies  year-to-year other  greater with  Population  of  stability  this  than any r e p o r t e d  previous  studies  shrub-land generally  and  open  consistent  Kluane  size  is  other  limited  habitat  I  forest with  home r a n g e  o£ a  Lake  with  previous  lower  than  i n most  chipmunk s p e c i e s ,  f o r the genus.  the central  mountains o f Oregon).  and  but the  i s s i m i l a r to that  found  Home r a n g e s i z e s a r e a t l e a s t 4 t i m e s  is  in different  Nevada, C a l i f o r n i a ;  and  at  of the population  chipmunk s p e c i e s .  conducted  the  chipmunks  o f chipmunks.  previous  in  least  choice  at  the fact  distant  that  closed  use comparison that  they  locations  Rocky Mountains  found to  by  Habitat  were  (Sierra  l n A l b e r t a ; and  Yukon c h i p m u n k s  prefer  canopy f o r e s t , which i s  the f i n d i n g s of previous  studies.  37 CHAPTER 3:  SPACE USE  AND  SOCIAL STRUCTURE IN  THE  LEAST CHIPMUNK  INTRODUCTION  the  This  chapter  use  of  minimus)  in  stability rodents and  and and  Moss,  limit  an  bar  tian,  compared  pennsylvanlcus  spacing  study  occurs  social  has  dominance structure Of  and  (Tamlas  chipmunk p o p u l a t i o n s  exhibit  to  least  populations  l n my  (Calhoun,  study  small  fluctuate  (Taltt  Clethzlonomys  area, widely  i n numbers  to  food,  s e t out  and  to  three  In a t e r r i t o r i a l  areas  from  i n what  These  territoriality  system, animals  others  (Healy,  are  have  excluded.  based  1967).  form  social types  space-related of  social  Tamlas.  I s t h e most fixed,  on  (Chris-  Three  types for  can  routes,  cyclic  territoriality,  avoidance.  use  travel  e x p l a i n both  i f , and  Watson  space  chipmunk a t K l u a n e L a k e .  types,  tive.  1962;  Explanations  stable populations  were c o n s i d e r e d :  encounter  mates,  areas.  to e s t a b l i s h  least  Wynne-Edwards,  shelter,  from c e r t a i n  1967)  i n the  1949;  the r e g u l a t i o n  Interactions affecting  have been used  which  of other  fluctuations  have e a c h been p r e v i o u s l y d o c u m e n t e d these  s t r u c t u r e on  chipmunks  of  l o n g been a s s o c i a t e d w i t h  access  spacing  social  multi-annual  example,  Social  Chitty,  of  1985).  spacing  behaviour  Thi3  marked  Mlcrotus  1970).  1950;  Yukon.  For  animal's  effects  population  1985).  i t completely  spacing  of  2)  population density  and  a  southwest  Wingate,  Social  or  the  in  which e x p e r i e n c e  rutilus  of  space  (Chapter  Krebs,  (Krebs  d i s c u s s e s the  restric-  defended  Definitions  core  of t e r -  38 ritoriality Pitelka,  vary, emphasizing  1959;  studies  of  Yahner  (1978)  Schoener,  Tamias  Broadbrooks  typical  and  f o r western  social  territorial.  territorial  suggested  behaviour i n  territoriality  was  to t e r r i t o r i a l i t y ,  i s lacking.  within  their  Numerous  b u t an  I n d i v i d u a l s do n o t but a r e dominant i n  c o r e a r e a s a n d have studies  on  prior  chipmunks ( e . g .  1978) have r e p o r t e d t h e e x i s t e n c e o f t h i s  system.  regular  avoidance Space  intervals  distance  I applied  to  animals  will  neighbours.  t o see the  would  which  least  core  of  areas.  but p r e d i c t s  these at  with A  does  chipmunks by by s p a c i n g a t  o r b y m a i n t e n a n c e o f a minimum  to detect  dominance  in  In t i m e , e i t h e r  Spacing  chipmunk  be s u f f i c i e n t  Space-related  documented  i s not a c t i v e l y  b u t by a v o i d i n g  have home r a n g e s  non-overlapping  first  a t a g i v e n time,  between  tested  was  is partitioned  by a g g r e s s i v e b e h a v i o u r ,  areas,  In  c h i p m u n k s were  and e x c l u d e a l l i n t r u d e r s ,  1970; E l l i o t t ,  (1981).  ranges  1939).  (1953), and  i s similar  core area  interactions  Encounter Getty  (Noble,  (e.g.  Yerger  of  Brand(1976)  access to resources there. Dunford,  these  occupancy  chipmunks.  e x c l u s i v e l y defended  agonistic  defense (1940),  evidence  dominance  territories  exclusive  or  Burt  that  (1970) f o u n d  Space-related  patrol  1968)  striatus,  claimed  Tamias amoenus,  either  maintained  interactions. 3  systems  Kluane  minimal  of  Lake.  overlap  space use Territorial  and m a i n t a i n  s y s t e m a t i c m o n i t o r i n g o f home this. not  aggressive defense  predict  exclusive  of core areas.  core  Encounters  39 should  result  in  agonistic  interactions  determined  by t h e l o c a t i o n  be  by o b s e r v i n g and comparing  tested  different  locations  Encounter frequent  of the encounter.  on t h e s t u d y  avoidance  than would  predicts  be e x p e c t e d  by  telemetry  comparing with  constructed patterns.  presence  radius social  of  real  define  chipmunks a r e the  encounter  using I  the  close  based  m.  home  from  t o each I n my  An e n c o u n t e r  This  range c a n be  e n c o u n t e r s measured by a  parameters as  be l e s s  o f home  ranges.  frequency  encounter  will  on t h e d e g r e e  of  of the other. 30  encounters  frequency  enough  p r e d i c t i o n can  outcomes o f i n t e r a c t i o n s a t  that  home r a n g e  an  This  dominance i s  site.  o v e r l a p and random movement w i t h i n tested  i n which  an  simulation  a n d random movement event  other that analysis  model  I  in  each used  which  two  I s aware o f an e n c o u n t e r  does n o t n e c e s s a r i l y  involve a  Interaction.  METHODS  The  study  described calculated the  ln  area,  Chapter  trapping 2.  t h e home r a n g e  Turner,  is  simple  to  telemetry  t e c h n i q u e s were  From t h e t e l e m e t r y a n d r e c a p t u r e d a t a I a r e a and o v e r l a p f o r each a n i m a l u s i n g  minimum c o n v e x p o l y g o n method  method  and  use,  1969) a n d f a c i l i t a t e s  ( S o u t h w o o d , 1966, p . 2 6 2 ) .  statistically comparison  stable  This  ( J e n n r i c h and  with previous  studies.  40  Behavioural  observation  During the rates  of  summer of 1985  behavioural  I i n v e s t i g a t e d the occurrence  interactions.  Population  density  and and  a c t i v i t y l e v e l s of chipmunks were too low f o r n a t u r a l i n t e r a c t i o n r a t e s to provide s u f f i c i e n t data to a s s e s s s p a c i n g therefore  provoked  interactions  seeds a t s i t e s spaced where chipmunk observed  i n t e r a c t i o n s among  a t t r a c t e d t o the seed  highest.  more of  them was  pelage-marked  an i n t e r a c t i o n ,  scored  of the  and  ended  when one  animal appeared. recorded  an  d i s p l a c e d the  loser  the seed  from  interaction p i l e by  such as t h r e a t s , lunges, chases been  described  (1979) f o r the e a s t e r n  chipmunk.  in  which  or For and  means of a g g r e s s i v e and  by Aniskowicz A draw  the winner  fights. and  occurred when animals  but n e i t h e r d i s p l a c e d the o t h e r .  o c c u r r e d when animals  showed no a g g r e s s i v e behaviour,  or ignored each o t h e r .  Similar  Vaillancourt  behaved a g g r e s s i v e l y  amicably  two  as e i t h e r a w i n - l o s s , a draw, or t o l e r a n c e .  d e f i n e d as  have  which were  A bout began when  p a r t i c i p a n t s were  A w i n - l o s s was  behaviours  chipmunks  d i s p l a c e d , or when a new  each bout the i d e n t i t i e s  behaviours  From a d i s t a n c e of 5 m I  were d i v i d e d i n t o bouts.  chipmunks began  was  sunflower  piles.  or more  the bout  p l a c i n g p i l e s of  I  s y s t e m a t i c a l l y over the g r i d s i n open areas  a c t i v i t y was  Observations  by  behaviour.  Tolerance but behaved  41 Encounter  avoidance  I n 1986 t e l e m e t r y was u s e d avoiding with  encounters.  encounter  within  sampling  using sampling  ranges  a n d compared  inter-neighbour counters.  time  (program  distances  locations  not consider t h i s  the  same g r i d  rarely  varied  than  a t t h e same  random model  with a c t u a l attributes  ln  encounter from  is  the  the telem-  m  l n length. of  are  o f 30 m. A l l  classified  l n encounters  radio-collars  reason  9.  four I used  In t h i s or  more  as en-  simultaneous  average  elapsed  i s 15 m i n u t e s ,  of f u n c t i o n i n g chipmunks  a t t h e same analysis animals  only the grid  I  on  grid  time  I used carried  H telemetry  radio-collars, X equipped  there with  time.  (which g e n e r a t e s encounter  d a t a t o compare  rates)  utilizes  the  following  actual  data.  For  each  simulated,  In each  a l l animals are  radius  Since the  0 to  which  4  of  t o be s e r i o u s .  from  the shortage more  transmitters  movement  30  of animals  For t h i s  Due t o  The  to  model  f r e q u e n c y from  d a t a do n o t p r o v i d e t r u l y  problem  windows  transmitters.  were  up  compared  i n Appendix  an e n c o u n t e r  number o f a n i m a l s w e a r i n g  only sampling  data.  against  were  listing  positions  estimates f o r a l l animals.  between  The  among  The t e l e m e t r y  positional  simulation  windows 90 m i n u t e s  window d i s t a n c e s  calculated  on  home  whether a n i m a l s  f r e q u e n c i e s were  by a  I obtained estimates of encounter  e t r y data  do  encounter  f r e q u e n c i e s generated  random movement 1).  Actual  t o determine  positions  of  nest  site  home  range  chipmunk  whose  and c e n t r e o f  42  activity model  were o b t a i n e d f r o m t h e t r a p p i n g  also  uses  values  for  activity,  time a c t i v e  above  location  that  derived  were  l o c a t i o n d a t a were mapped study grids The 3.1.  algorithm  Initial  sampled  and  whether  or  leaves  its  randomly  input  not t o nest  chosen  chipmunks  beyond  this  chosen  that  process  repeats  and  moves any  the  the  radius  to determine  simulates for  distances  one  the p e r i o d  be compared  day's that  s y s t e m as  identity  the  in figure  of  of  360  chipmunks  (15  maximum  and  a new  positions  time in  distance  of s i m u l a t e d  activity.  The  In a  randomly  distance.  interval,  t h e model  This  when t h e  are  encounters.  from  movements  direction  w i t h the s p e c i f i e d  t h e number  m)  Simulated  90 m f o r f e m a l e s ) ;  each  a r e compared  i t moves i t  degrees.  i t s maximum  of  corresponds to  if  coordinate  specified  end  d i s t a n c e s among a l l chipmunk These  Computer g e n e r a t e d  i s 50%).  1 grid  the animal w i t h i n  lated.  of  of changing  is Illustrated  t h e number and  a r e preempted,  until  probability  t h e model  (180 m f o r m a l e s ,  boundary  The  from c e n t r e  same c o o r d i n a t e  (probability  within  of a c t i v i t y  keeps  the  direction:  centre  and  data.  When t h e r u n b e g i n s , e a c h a n i m a l d e c i d e s  move  remain  distance  from the d a t a .  in  includes  site  telemetry  comparison.  used  t h e month.  maximum  ground,  on  to f a c i l i t a t e  and  calcu-  encounter Each  run  number o f e n c o u n t e r s g e n e r a t e d the sampling  window c a n  w i t h number o f e n c o u n t e r s r e c o r d e d i n t h e  field.  then  43  Flowchart of random-encounter s i m u l a t i o n model.  # of  stop  chipmunks  chipmunk identities Define:  -naat alt«« -aotlvlty centra -max. rang*  set time  start day  choose direction calc. new location  number of encounters  month  i length of active day  start from nest sites  t  snapshot  encounter  interval  radius  encounter statistics relocate —  1  45  RESULTS  Home range o v e r l a p  Home  range  overlap  measuring the percentage I calculated data.  between of  neighbours  spatial  overlap  polygons.  o v e r l a p s e p a r a t e l y u s i n g the t r a p p i n g and t e l e m e t r y  recaptures  overlap.  were  Included  Five captures  e s t i m a t e s of  i n c a l c u l a t i o n of home range  i s seldom  sufficient  crowding  to  yield  accurate  home range a r e a , but i n c l u d i n g these home ranges i n  the o v e r l a p a n a l y s i s g i v e s a b e t t e r  i n d i c a t i o n of  the degree of  than c o u l d be o b t a i n e d by e x c l u d i n g them.  According to  trap-based e s t i m a t e s  of home range,  mean home  combined.  Each home  of 14.9 o t h e r s  (Table 3.1).  range o v e r l a p was 94.3% f o r a l l animals range was  overlapped  O v e r l a p on female does  by  home  not d i f f e r  an average  ranges  by  Male home ranges  neighbours  female  neighbours  than  other females  significantly  o v e r l a p , 70.4 %.  sex  of convex  by  For the t r a p p i n g d a t a , a l l chipmunks with more than 5 non-  linear  this  was determined  home  were  ranges  averaged  75.1%;  from male-male home range overlapped are  by  more male  overlapped  by female  (6.5 v s . 4 . 0 ) .  T h i s i s c o n s i s t e n t with the male-biased  telemetry-based  estimate of home range o v e r l a p was 86.8%  ratio. The  46 for  a l l Individuals  since use,  not  a l l animals  area  an  than  average  exclusive  a r e a s , as its  a core  area  do  In  terms  overlap revealed  86.8%.  not,  by t e l e m e t r y  a chipmunk  any  neighbouring  therefore,  frequents  home r a n g e maps a r e p r e s e n t e d  home r a n g e  home  was  range,  home  range  radio-locations.  correspond  to core  t o be t h e p a r t o f  t h e most.  in Figures  estimate,  of i n t e n s i t y of  Within  Is g e n e r a l l y u n d e r s t o o d  the animal  3.1—Mean  the trap-based  o f o n l y 8% o f t h e " o w n e r ' s "  areas  range which  Table  than  which does not o v e r l a p with  contains These  higher  lower  were r a d i o e d .  t h e amount o f home r a n g e  substantially the  combined,  Telemetry-based  3.2 a n d 3.3.  overlap  SOURCE  PERIOD  Trap  2.5  mo.  94.3  75.1  70.4  14.9  4.0  6.5  Telem  4.5  mo.  86.8  65.2  82.0  6*  3*  5.2*  Telem  12 May10 J u n e  80.1  60.0  73.6  5*  2.5*  4.7*  Telem  11 J u n 20 J u l y  76.2  52.0  76.0  5*  2.5*  5*  Telem  21 J u l 12 S e p t  83.0  62.6  78.8  6*  3*  5*  Home September, weaning data  range  shifts  especially  their  into  PERCENT OVERLAP ALL F M  three  young.  occurred among  between  females  I therefore  p e r i o d s based  NO. NEIGHBOURS OVERLAPPED ALL F M  that  divided  on f e m a l e  May  and  had all  breeding  just of  t h e end o f finished  the telemetry  condition:  pre-  47  Fig.  3.2  —Telemetry-based  f r o m 1 t o 37 on the  grid  home r a n g e  t h e y - a x l s and  boundaries.  One  grid  from  map  1-29  of G r i d on  coordlnate=15  H..  Coordinates  the x - a x i s are m.  within  49  Fig.  3.3  --Telemetry-based  f r o m 1 t o 29 coordlnate=15  home r a n g e  on b o t h a x e s a r e w i t h i n m.  map grid  of G r i d  X.  boundaries.  Coordinates One  grid  X  51 lactating  (May 12 t o J u n e  post-lactating less  during  entire most  these  field often  greater  (July  21 t o September  discrete  season,  near  their  nest  range o v e r l a p i s  when c a l c u l a t e d  lactating  observations during  mating  season  and  chipmunks,  14 o f them on g r i d  240  of observation,  hours  were a g g r e s s i v e .  an a v e r a g e  of twenty  interacted  interactions 353  Home  over the  females,  sites,  20) a n d  overlap  who  were  i s always  observation  the  animal  11 t o J u l y  t h a n 50%.  Behavioural  (89%)  for  (June  12).  p e r i o d s than  b u t even  located  Behavioural  after  10), l a c t a t i n g  bouts  Each bout  would  yield  The  data  dominance  1985 b e g a n  X.  on 35 During  o f w h i c h 315  animals,  so t h a t  f o r each a n i m a l .  of 6 neighbours.  Bach  Thus, even i f  among n e i g h b o u r s ,  the t o t a l of  only  per  obtained  reversals  21 on g r i d  i n v o l v e d two  an average  of  were r e c o r d e d ,  were r e c o r d e d  evenly  summer  i n t e r a c t i o n data  H, t h e o t h e r  divided  recorded  neighbours. document  were  provided  353 b o u t s  bouts  with  the  for  are each  3.3  bouts  therefore  pair of  too sparse to  individual  with  its  neighbours. However, t h e r e l a t i o n s h i p be all  between d o m i n a n c e and l o c a t i o n c a n  e x p l a i n e d by l o o k i n g a t t h e d a t a win-loss  bouts,  w i n n e r s and l o s e r s  there  l n the  which the i n t e r a c t i o n  on a b o u t - b y - b o u t  was no s i g n i f i c a n t mean  occurred  distance  from  to the centre  basis.  difference the  For  between  location at  of the p a r t i c i p a n t ' s  home r a n g e .19,  df  on e i t h e r  = 109;  162).  nest  was  site  from both  (grid  X, p a i r e d  H,  paired  samples  grid  Nor was  analysis  grid  there  repeated  (nest  sites  any  significant  for  distance  were  grids, paired  samples t t =  = 1.32,  .41, p = .68, d f =  difference  from  known f o r 19  p =  when t h e same  interaction location to chipmunks).  Using  s a m p l e s t = .67, p =.51, d f = 220  data (Table  3.2).  Table  3.2—Mean  GRID  distances  from  CATEGORY  Interaction  t o home  MEAN DISTANCE WINNERS  IN METERS (SD) LOSERS  X  home=centre home=nest  87.6(83.75) 91.1(57.32)  79.2(49.61) 95.1(44.72)  H  home=centre home=nest  72.7(53.18) 85.1(55.71)  75.8(51.00) 89.3(48.11)  Only  bouts  between a n i m a l s  encounter  occurred  estimates  o f burrow s i t e s  for  non-residents.  between r e s i d e n t s these  resulted  instances The  were  differs  and  were  as  to  7.0,  p < 0.01, d f  bouts  that  were  intruders.  over  frequency  the  unavailable occurred  Bouts  with  the only  entire This  expected  i n t e r a c t i o n s as  since  Twenty-four of  t h e 28 e n c o u n t e r s .  from the  1).  analysis,  (8% of the t o t a l )  I witnessed  7 of  where t h e  i n t e r a c t i o n s and i n c l u d e d  dominate i n =  the g r i d  i n the previous  non-resident  won o n l y  on  a n d home r a n g e c e n t r e s  from unprovoked  significantly likely  included  Twenty-eight  of f i g h t i n g  intruders  resident  season.  frequency  i f intruders  residents  Intruders  were  (X * = s  usually  53 initiated on  by  residents  the g r i d .  Of t h e b o u t s  non-breeding  animals  For a l l was s l i g h t l y losers  were s u b o r d i n a t e t o o t h e r  lost  by i n t r u d e r s ,  w h i c h were  win-loss bouts,  lighter  higher  to determine  whether s e x ,  t o dominance.  t h e outcome o f i n t e r a c t i o n s  Adults  won  in a l l  not  in  more .001,  with other adult  than  d f = 2 ) . Bouts  3 categories  May  than pregnant  females  involving  Lactating  breeding  condition  20,  females  females  =  males  (X  a  = 15.51, p <  were d i v i d e d  (X =5.12,  were  significantly  interactions a  a  most b e h a v i o u r a l  won  bred  (X  of breeding  t h e y were pregnant,' won  = 1).  a g g r e s s i v e than  a t which p o i n t  breeding females  females  d i d not  .40, d f  The e f f e c t s since  of  tests  with juveniles  which never  and p o s t - l a c t a t i n g  weight  Chi-square  t o be more  Breeding  a c c o r d i n g t o whether  or p o s t - l a c t a t i n g .  while  tended  place after  interactions  If  mean  .72, p =  residents.  breeding condition.  Encounter  =  were a n a l y z e d o n l y f o r f e m a l e s , took  (50.9 g)  of the p a r t i c i p a n t s a  to  Intruders.  the  aggressive interactions  19.37, p < .001, d f = 1) w h i c h interactions  (X  lost  of winners  a g e , and  The s e x  affect  observation  the  than  residents  71% were  g, t = 3.43, p = .001, d f = 3 1 4 ) .  were r e l a t e d  condition  than  t h e mean w e i g h t  but s i g n i f i c a n t l y  (49.3  were u s e d  that  into  lactating, more  p<.025,  often df=l).  avoidance  encounter females  avoidance  are  occurs,  lactating,  since  i t should their  be most  apparent  movements  a r e more  54 r e s t r i c t e d and they win a g r e a t e r  p r o p o r t i o n of  t h a t time.  frequencies during p r e - l a c t a t -  Therefore,  encounter  interactions at  i n g , l a c t a t i n g , and p o s t - l a c t a t i n g p e r i o d s were compared s e p a r a t e l y with 3.3.  For  frequency  the model's output. the  pre-lactating  and  was  significantly  less  random model, so df=45).  the samples  For the  lactating than  were  that  pooled  3.3—Mean encounter  from the model  frequency on  i n Table  p e r i o d s , encounter generated  (t=2.41,  p o s t - l a c t a t i n g p e r i o d , encounter  not d i f f e r s i g n i f i c a n t l y  Table  The r e s u l t s a r e presented  p  by the = .02,  frequency d i d  (t=1.30, p=0.20, df=42).  g r i d H vs. s i m u l a t i o n  PERIOD  MEAN ENCOUNTERS  Pre-lactating  0.48  1.16  Lactating  0.86  1.19  Post-lactating  1.37  1.23  MEAN ENCOUNTERS GENERATED  DISCUSSION  From the home range data presented here, the h y p o t h e s i s t h a t l e a s t chipmunks a t Kluane ly  rejected.  range  Both  Lake a r e t e r r i t o r i a l can be c o n c l u s i v e -  t r a p and telemetry-based estimates of home  o v e r l a p demonstrate  that e x c l u s i v e core areas do not e x i s t .  55  T e r r i t o r i a l i t y can a l s o be r e j e c t e d u s i n g Noble's (1939) d e f i n i t i o n of  territory,  c l u s i v i t y as  which  emphasizes  d i a g n o s t i c of  territoriality.  l a r g e t o be p a t r o l l e d and defended. the same  g r i d areas  these a r e a s ; The  defense  for a l l  I n t e n s i v e use i s c e n t r e d on  they cannot be defending  core  l n chipmunks  Tamlas s t r l a t u s  has  never  have  considered  o n l y l n order  to evaluate  species  of  western  evidence  S o c i a l dominance  been  studies.  f o r dense The s o c i a l  and  breeding  importance (Heller,  of  to  was  found  appears to  previous chipmunks among  1971; Sheppard, 1971;  for space-related  dominance.  be based on I n d i v i d u a l c h a r a c t e r i s space.  Size,  c o n d i t i o n a r e t y p i c a l l y r e l a t e d t o dominance l n  dominance was e v i d e n t  Non-residents  component.  An  space-independent p a t t e r n of s o c i a l  i n the I n t e r a c t i o n s  r e s i d e n t s which d e f e a t e d r e p o r t e d above.  system of the  1978).  the a p p a r e n t l y  non-residents.  p o p u l a t i o n s of  competition  s t u d i e s of s o c i a l dominance t h a t l a c k any s p a t i a l exception  Territorial-  examined,  t i c s such as s i z e and age r a t h e r than ownership of age  a t Kluane Lake  a g o n i s t i c behaviour i n l e a s t the  overlap  areas.  specifically  chipmunks  S t a t e s , 1976; C h a p p e l l , Little  reported  and Tamlas amoenus.  l e a s t chipmunk has studies  been  home ranges  l n Tamlas minimus  does not s p e c i f i c a l l y c o n t r a d i c t p r e v i o u s ity  than ex-  Home ranges a r e too  animals whose  absence t e r r i t o r i a l i t y  rather  were  them, which  between r e s i d e n t s and  g e n e r a l l y heavier i s contrary  to  than the the t r e n d  Three of the i n t r u d e r s were r e s i d e n t on another  g r i d where each of  them  was  socially  dominant.  These  three  57 total. There agonistic  is  social  n e s t s were group  factor  structure  within  15  which c a s t s doubt  revealed here.  m of another  male and  of August.  nests,  yet  Brown  (1964)  female These  chipmunk  suggested  that  economically defendable.  a  rich,  of  which  chipmunks  were f r e q u e n t l y o b s e r v e d dia  berries  at  meters without  any  sign  distances  to  aggressive  studies provoked social  However, t h e provoked  of  chipmunk  maintained  chipmunks,  absent  relationship  interactions  the  grids.  is  only  so t h a t i t constituted  of  presence the  1986,  Shepher-  than  In 1985,  piles  were o f t e n  social  behaviour  normal  chipmunks  abundant  d i s t a n c e s of l e s s  m a t seed  the  their  typical  of a g g r e s s i v e behaviour.  interactions.  s p a c i n g was  piles  In August  as  three inter-  sufficient  interactions.  aggressively of western  Kluane.  on  behaviour  and  a  while sharing  piles  f e e d i n g on s e a s o n a l l y  of 7  Previous studies reported  not  inter-individual  individual provoke  at  were  were weaned a t  seed  resource,  located  a nest,  is distributed  sunflower  defendable  in behaviours  behaviours  My  sharing  a t seed  resource  the  In a d d i t i o n ,  amicably  territorial  the c o n t e s t e d  discrete,  behaved  of  nest.  juveniles  aggressively  is  resulted  after  animals  interacted  a d a p t i v e where  pair  on t h e a p p a r e n t l y  Six  o f t h r e e m a l e s were r e p e a t e d l y o b s e r v e d  were a end  another  and  s p a c i n g mechanisms,  have r e l i e d  I believe a t Kluane between naturally  that  either  reasons  frequency of  occurring  have  especially  lab t r i a l s  aggressively  f o r the the  on  that  or  maintained  given  above.  artificially  interactions  needs  58  to  be s t u d i e d On  the  basis  the  data  nce  occurred.  data  before  this  of  conclusions  the d i f f e r e n c e  and t h e random m o d e l , The  encounter  radius.  provided  simultaneous data,  average  chipmunk  home  constraints  such  tion,  the  and  resources, that  and  occurring Kluane.  the  restriction  from  as  that the of the  would  large  forested  have as an  habitat  among  of  adults  that  rare  encounter  of low-  by e c o l o g i c a l  movements d u r i n g  lacta-  seasonally-abundant Interactions.  Mutual  was r e l a t i v e l y  structure  i s determined  chipmunks  of August.  suggests  avoida-  length  sampling  the  post-lactating period  of nests  aggression  by t h e  social  space-related  resulted  the f a c t  area  with  Tamlas minimus  the  t h e month  This  that  than by  in  the sharing  and  n e c e s s i t y of harvesting  random may have in  of  f o r an  encounter  by  scan  rates in  was n o t f e a s i b l e .  suggest  as the  rather  spacing  berries  range,  s i g n a l bounce, t h i s  density populations  and  Automatic but  that  limited  simultaneously,  defined  These r e s u l t s  is  confirmed.  between e n c o u n t e r  I conclude  comparison  were n o t g a t h e r e d  causing  c a n be  The f a c t  d i d not d i f f e r  harvesting tolerance  indicate  food  from  Shephezdla  while  feeding  that n a t u r a l l y  among l e a s t avoidance  chipmunks a t i s maintained  passively. It  is  also  possible  behaviour  occurred  other.  Relatively  longevity  of  1943)  could  observed  among c h i p m u n k s high  the species result  that  in  which  survival (up t o  a high  rates  6 years  degree  instances were  of amicable  related  (Chapter according  of relatedness  t o each  2) a n d t h e to C r i d d l e , among  neigh-  59 bours. Heller were  less  although  (1971) and aggressive  this  (Sheppard,  than  1971;  to  the  the  have been  least  chipmunks  species  of  their  i n the  constant, Aggressive  ted  the  during  the  suggests  short snow-free  least  Aggressive  Isolated  metabolic  requirement  that  occupies.  Yukon a r e  near  the  in  chipmunk  behaviour and  from  congeneric limit  may  in  sufficient  fact food  may  aggresspecies, of  their  T h e y may  very  due  sage-brush  larger  northern  of  aggressive  spacing behaviour  of r e s o u r c e s  season.  concept  the  constraints apply.  of hoarding  congeners  with  p o p u l a t i o n d e n s i t y , and  defense  the  ensue  by c o e x i s t e n c e  low,  same  invokes  i n the  f o r a g g r e s s i v e l y maintained  ranges. by  and  the  t h a t would  congeners.  r a n g e where d i f f e r e n t need  (Brown, 1964)  inhibited  dominant  little  Heller  chipmunks  in adjacent habitat,  s t u d i e s of  1978).  stress  that least  living  metabolIcally infeaslble  habitat  sively  congeners  Chappell,  physiological  desert also  is  (1976) f o u n d  c o n t r a d i c t e d other  economic d e f e n d a b i l i t y behaviour  States  have  because  l a r g e home be  prohibi-  for  winter  60 CHAPTER 4:  THE INFLUENCE OF GRID-TRAPPING ON SPACE USE AND HOME RANGE  IN THE LEAST CHIPMUNK  INTRODUCTION  This trapping a  chapter  on home r a n g e .  trapping  that  Investigates  grid  live  availability  o f two  changes s u f f i c i e n t trapping an  affects  accurate  Firstly,  changes  there.  two  possible  presence  resources: to distort  home r a n g e s of  food  traps and  home r a n g e ?  home r a n g e  of the animals  could  increase the  cover.  Are these  Secondly,  t h e measurement o f home r a n g e .  measure o f  of g r i d -  i t a s k s whether t h e p r e s e n c e of  the a c t u a l  The  effects  compared  i t asks  Does  to that  how  i t provide afforded  by  telemetry? Criticisms estimation 1964; are of  of  are  grid-trapping  common  as  (Hayne,  1950;  Banks e t a l , 1975; C r a n f o r d , confined  to properties  recaptures,  cient and  lack  by  grid-trapping  contain McMahon,  because  t h e movements o f 1986).  measurement  These  o f home r a n g e  f o r home  range  Sanderson and Sanderson,  1977), but p o i n t s  required  assumptions about  of continuous data.  method  of c r i t i c i s m  o f g r i d - t r a p p i n g s u c h a s l o w numbers  t h e amount o f t i m e  captures,  a  travel  Home r a n g e grids  are  animals  that  aspects  of  rather  to  accumulate  distance i s often  suffi-  between  traps  underestimated  not s u f f i c i e n t l y i n h a b i t them  large to  (Broome and  grid-trapping  affect  t h a n t h e home r a n g e  itself.  the  61 Previous imental et  studies  1983;  al,  employed studies affect  studies  would  to  be  the very own  parameter  observations leaving  enough  or  food  movement p a t t e r n s . one  pouches with as  their  I  bait  sessions,  and found  chipmunks t o food  observed  another  that  piles  along  both  t o measure.  sessions  animals  a grid  during  i t was  placed  of such  methodologies  the chipmunks' environment  filled)  Sullivan generally  The v a l u e  between  and b e d d i n g , and p a u s i n g  pouches  exper-  t h e summer o f 1 9 8 5 s u g g e s t  repeatedly  trap after  have  i f their  pre-balted  to  1982;  1985),  range.  are intended  from  traps  cover  home  compromised  they  especially  Mares e t a l . ,  Trombulak,  estimate  severely  t r a p p i n g and  (visiting  1984;  Lack! e t a l ,  home r a n g e ,  al.,1976;  (Mares e t  trapping  My  o£ chipmunk  "trap-lining" cheek  to scatter-hoard  and  generally  added  to affect  row, f i l l i n g  only  that  between . t r a p p i n g easier  a t t r a p - s t a t i o n s than  to attract elsewhere.  Predictions  Providing generally tive  thought  studies  availability  1975),  supplemental  and  reductions  food  to decrease  have  shown  a  negative  mammal  et a l . ,  to small  scale  food  is  revealed  home r a n g e a r e a s  should  correla-  decrease  1981).  addition, after  food  eta l . ,  have r e s u l t e d i n home  Taitt  trapping  Numerous  1943; O'Farrell  1976;  (Mares e t a l . ,  home r a n g e s i s  r e l a t i o n s h i p between  (Blair,  a d d i t i o n experiments  equivalent  small  home r a n g e a r e a .  a n d home r a n g e s i z e food  to  range  If gridtelemetry-  grid-trapping is  62 Instituted  on a n a r e a ,  Another expansion that  possible  trapping  that  w i t h a change  begun, a n i m a l s of their  Intensively occurring,  to  periphery  s t a t i o n s more  part  frequently  should  with  This  traps  predicts  inward  should  respond  from  use t h e  does not.  the g r i d  food-enriched  the  grid,  more  If trap-liningi s  should  than would  to grid-  Once t r a p p i n g h a s  which o v e r l a p s  which  inside  area  distribution.  home r a n g e , t h e p a r t  then animals  i s a s h i f t or  supplied  home r a n g e s .  ceases.  i s operating.  the g r i d  i n use  the  areas  those  the g r i d  on  on t h e g r i d  than  grid-trapping  g r i d s a n d home r a n g e s h i f t s  while  live  after grid-trapping  include  within  onto t r a p p i n g  Animals  trap  to  were n o t p r e v i o u s l y  edge o f t h e g r i d  part  response  o f home r a n g e s  immigration the  and i n c r e a s e  be  be e x p e c t e d  located  near  by c h a n c e .  METHODS  Procedures  used  i n the c o l l e c t i o n  data  have been d e s c r i b e d  will  n o t be r e p e a t e d  Experimental  In A p r i l were s u r v e y e d grid On  H  July  (the  extensively  of trapping in  previous  and  telemetry  c h a p t e r s and  here.  design  1986 two in similar southernmost  18 ha  grids, approximately  habitat. grid)  16, I removed a l l t r a p s  From e a r l y  was o p e r a t e d from g r i d  H  1 km  apart,  May t o m i d - J u l y ,  as a t r a p p i n g and t r a n s f e r r e d  grid. them  63 to g r i d  X (the n o r t h e r n g r i d ) .  g r i d X alone was t r e a t e d as 2.1).  From m i d - J u l y to l a t e  a trapping  grid  (Figure  September, 2.1, T a b l e  I used r a d i o - t e l e m e t r y to monitor home ranges of r e s i d e n t  a d u l t s throughout the summer. On each g r i d , t r a p p i n g range s i z e s  as a  r e v e r s e d on g r i d  The sequence  of treatment and c o n t r o l were  X with r e s p e c t to g r i d H l n order to account f o r  effects.  according to  Chipmunk  home  ranges  Broadbrooks,  1970).  If  home ranges  while for g r i d X  residents  seasonal e f f e c t s  determine  size, grid  in early  ranges  movements d u r i n g  the  reverse  should  home range s i z e  Home  e f f e c t s on food  i n l a t e summer, be  l a c t a t i o n would  true.  If  then s e a s o n a l changes  f o r both g r i d s .  i n l a t e than i n e a r l y summer on both  Home range  to vary  H residents should  summer than  i n home range s i z e s h o u l d be c o n s i s t e n t t i o n of  known  l a c t a t i o n (Martinsen,  grid-trapping  a v a i l a b i l i t y determine home range have s m a l l e r  are  season, e s p e c i a l l y f o r females, whose movements a r e  r e s t r i c t e d d u r i n g l a t e pregnancy and e a r l y 1968;  treatment, and home  d u r i n g the non-trapping p e r i o d were taken as c o n t r o l  values f o r that g r i d .  seasonal  was used  result  Restric-  i n l a r g e r home  grids.  calculations  range  polygon method minimum convex  size  was  calculated  (Southwood, 1966). polygon  reaches an asymptote  It  u s i n g the minimum convex i s well-known  that the  (MCP) i n c r e a s e s with sample s i z e u n t i l i t  a t the a c t u a l home range s i z e  (Stlckel,1954;  64 J e n n r i c h and the  Turner,  e a s t e r n chipmunk,  captures.  Therefore,  ments t o sample for  a l l  size,  an average  after  20  estimates  because  MCP  of  collect not  Effect  In the  In  trapping  to  estimates  are  same p e r i o d  only  range  estimate  used.  home  I  range range  of locations.  p o i n t s i n each  problems  i n some  For  I could  cases.  only  I chose  estimate  trap-based MCP as for  with  or l a t e  is  such  the a n a l y s i s .  used  exist  i s revealed  f o r the  20 l o c a t i o n  to l o g i s t i c  on home r a n g e  compared  (early  home  of locations  compare  is  in this  i n s t e a d o f a t 9 0 % home  more t h a n  estimates,  points data  from  source  telemetry-based location  (1980)  b u t due  these  order  95%  captures  t h e minimum number  locations  chipmunks range  t a r g e t f o r minimum number  t h e summer,  of data  o f home  measure-  m a j o r i t y of s t u d i e s f o r which  9 0 % or  I collected  that f o r  o f f a f t e r 20  a t 20 r a d i o  radio-collared  8 7 . 1 8 % (SD=13.96)  20  showed  levelled  MCP c a l c u l a t e d  Mares e t a l .  to exclude  area  (1980)  t o s t a n d a r d i z e home r a n g e  For a l l  after  a field  most a n i m a l s  range  I used  are calculated,  I used  as  of  Mares e t a l .  In o r d e r  locations.  calculate  limit  home  comparisons.  study  part  1969).  home r a n g e e s t i m a t e s  calculated above. half  Since, of  the  telemetry data  summer).  using  the  with  first  20  f o r each  chipmunk,  summer,  trap-based  collected  during the  RESULTS  Effects  of g r i d - t r a p p i n g on home range  Table 4.1 p r e s e n t s the mean t e l e m e t r y - b a s e d for early  and l a t e  summer on  listed  i n T a b l e 4.1  range s i z e s opposite  to  that  predicted  e f f e c t , but t h i s t r e n d  home range  g r i d H and g r i d X. suggest a  trend  size  The mean home i n home range  f o r a g r i d - t r a p p i n g food  addition  i s not s t a t i s t i c a l l y s i g n i f i c a n t :  grid H  p a i r e d samples t=.37, p=.72; g r i d X p a i r e d samples t=.41, p=.69.  Table 4 . 1 — T e l e m e t r y - b a s e d home range e s t i m a t e s . GRID  EARLY SUMMER MEAN(N) S.D.  LATE SUMMER S • D. MEAN(N)  H  2.8 ha (11)  1.9  2.7 ha (11)  1 .6  X  2.2 ha (8)  0.9  2.4 ha (8)  1 .0  There  i s no  detectable  food a d d i t i o n e f f e c t due t o g r i d -  t r a p p i n g on the amount of immigration o c c u r r i n g on the two g r i d s . Three  adult  chipmunks  immigrated  onto  summer while I was t r a p p i n g on g r i d H. was t r a p p i n g adult  on g r i d  X, 2  In  each g r i d d u r i n g l a t e summer,  a d u l t s immigrated  immigrated onto g r i d H.  early  while I  onto g r i d X and 1  Measurement of immigration was not  66 as  reliable  grid,  f o r the non-trapping  since  sightings  i t was  relationship recorded on  grid  X.  50%  number early  locations  summer, w h i l e  i n late  number  a  is  df=l,  telemetry within  Locations  classified  I  with  that  summer and more  (X2=16.7, regime.  df=l, On g r i d  I  can  of the  trap  H, t h e  number  in  l o c a t i o n s exceed the X, t h e  not  trend i s  effect  since  statistically  sig-  station  trend  as  o f 7.5  discern  "trap-associated",  that  must  be c l o s e  a chipmunk  For g r i d  less  frequently  This Is  X,  t h e same,  system.  enough t o  i s at least  locations are  than expected i n  expected  coincides  i t i s the  on t h e g r i d  trap.  than  s t a t i o n as "not  m because  based  assume  frequently  the  I classified a l l  7.5 m f r o m a t r a p  stations  p<.001). H,  On g r i d  expected  the grids  as " t r a p - a s s o c i a t e d "  trap  the  On g r i d  is  inside  than  could  aware o f t h e p r e s e n c e associated  i t  I chose a d i s t a n c e  distance  r a d i o - c o l l a r s , 10%  with a grid-trapping  but  of a  locations greater  trap-station  p<.01).  grid.  p>.10).  7.5 m  smallest  early  df=l,  on e a c h  the g r i d s .  summer, o f f - g r i d  locations  trap-associated".  with  i s l e s s than  causal  H and 3 s i g h t i n g s  2 sightings  outside  reversed,  a  were  of a  of  I n e a r l y summer I  on g r i d  equipped  i s consistent  (X =2.5,  locations all  (X =7.0,  which  treatment  For  a  were  locations  reversed,  and  summer t h e r e  trapping  The number  the absence  immigration.  o f unmarked a d u l t s  of o f f - g r i d  nificant  and  9 o f t h e 24 c h i p m u n k s of  expected  the  In l a t e  f o r the  systematically.  corroborates  between t r a p p i n g  5 sightings  For to  not monitored  o f unmarked a d u l t s  a 3 I t was  grid  in  with  late the  although  summer  trapping  i t  i s not  67 significant the  trapping  Effects  data  source  both  t h e same p a r t  d i f f e r e n c e between mean  from  revealed  This  trapping  home r a n g e  does  of that  on g r i d  with  X.  trapping  home  range  size  by t e l e m e t r y .  i s 2 . 4 h a , mean t e l e m e t r y t=1.63,  and  telemetry  o f t h e summer, t h e r e  and t h a t d e r i v e d  3.0 ha ( p a i r e d s a m p l e s  not c o i n c i d e  range  f o r which  available for  significant  p>.10).  on home  t h e 19 a n i m a l s  are  derived  df = l ,  r e g i m e , w h i c h was t h e r e v e r s e  of data  For  is  (X2=2.95,  revealed  i s no  ( a t n=20) Mean t r a p home  range  p=.121).  DISCUSSION  Effects  of g r i d - t r a p p i n g  Grid-trapping chipmunk  home  traps  does  not  ranges.  trap-ses3ions baited  on home  as  and  i n 1985,  cotton  trapping  certainly  increased  material,  and c o v e r ,  but  estimates  of  indicates  that  grid-trapping  home r a n g e  not that  f o r experimental provides  measurably  change  Chipmunks c o n t i n u e d  they d i d  of bait  range  were  to  independent  studies  an adequate  affect  of  estimate  of  during  t o empty p r e -  trap-sessions.  the a v a i l a b i l i t y enough  size  to trap-line  and c o n t i n u e d  between  the  of food,  Gridbedding  telemetry-based  of trapping.  chipmunk  home  o f home r a n g e .  This range, There  68 Is no e v i d e n c e  that  that  would  food  and d e n s i t y  invalidate conclusions  Another  of  onto the g r i d Immigration  In  late  response  food  occurred  al.,  increase  1976;  and  was  Indistinguishable  from  on t h e amount o f  immigration.  Tests trapping  f o r changes  were  home r a n g e s patterns  than g r i d s on  one g r i d  than  lie partially  so  that  only.  of  seasonally  inconsistency  or absence  females abundant  after  which  their  The t r e n d  of g r i d - t r a p p i n g ,  of g r i d -  chipmunks  are being  suggests  than  immigra-  result  t h e g r i d s change  trapped.  early  grids  G r i d - t r a p p i n g had  as a  grids that  1981).  on b o t h in  In  recruitment.  outside  frequent  This  Krebs,  i n the d e n s i t y of  I t i s possible that  that are not being  the presence  shifts  they  emergence  i n use d i s t r i b u t i o n  inconclusive.  and  an i n c r e a s e  tion  effect  by i m m i g r a t i o n  o f a d u l t s was h i g h e r  there  increase the  t h e same r a t e  on b o t h g r i d s a t j u v e n i l e  no  i n a way  range r e s p o n s e t o  caused  Gilbert  juveniles was  home  experiments which  a t approximately  Immigration  summer,  to  i s a density  (Mares e t  the season.  about  home r a n g e  manipulations.  typical  availability  over  grid-trapping distorts  whose  movement  trapped  more  is significant  that  factors  other  s u c h a s home  range  j u v e n i l e emergence or a v a i l a b i l i t y o f  patches  of food,  may  explain d i s t r i b u t i o n of  use. For of g r i d  radio-locations X show a  trapping. grid  This  change suggests  X b u t n o t on g r i d  H.  i n s i d e the in  use  that  g r i d s , only  concentration  trap-lining  Since  grid  X  the inhabitants in  behaviour contained  response t o occurred a  on  greater  69 proportion possible  of  greater grids  resource  mean than  over  from their  data  entire yield  higher  estimates  of data  Trapping  polygon  and  source  method  are  Sherman  range  of  o f 4.86  I reported  was  was a more  and  most  This  3 o f t h e male  are  before  therefore  lactating  y o u n g had  2 to calculate p r e - and  2.  d i e d or d i s p e r s e d  Secondly, their  ha f o r b o t h  i n Chapter  Firstly,  switched  females  emerged.  mean home r a n g e  Since f o r the  post-emergence p e r i o d s ,  home r a n g e  than  the  data  they  for either  on home r a n g e  and t e l e m e t r y p r o v i d e  estimates  sizes  both  X  is  4.1 a r e s l i g h t l y  l n Table  home r a n g e s  after  grid  2), It  alone.  Effects  able  Include  listed  range s i z e  analysis.  chapter  (Chapter  t r a p s on  o f two f a c t o r s .  home r a n g e s in  H  H.  which  treatment  this  used  summer  period  season  the largest  grid-trapping  shifted the  t h e mean home  i s the r e s u l t  chipmunks w i t h  excluded  sizes  grid  by  on g r i d  home r a n g e half  than  provided  than  the e n t i r e  discrepancy  the  habitat  t h a t the food  significant The  poor  of  home  o f home  equal.  range  range  This  home r a n g e e s t i m a t e s  and  size  using  calculation,  i s c o n s i s t e n t with  (1983) who compared  calculation  statistically  showed  eight  method.  t h e minimum c o n v e x provided  t h e sample  the f i n d i n g s  different  methods  of Jones o f home  t h a t t r a p - and t e l e m e t r y - r e v e a l e d  were most c o m p a r a b l e  minimum c o n v e x p o l y g o n  indistinguish-  t o one a n o t h e r  with the  70  At very  low sample s i z e s , t r a p - and t e l e m e t r y - r e v e a l e d home  range e s t i m a t e s observed  would d i f f e r ,  (Hayne,  because as  1949; S t i c k e l ,  c a p t u r e s o f t e n occur  numerous authors  1954; Jones and Sherman, 1983),  i n a straight  l i n e due  t o the  t r a p s , y i e l d i n g MCP home range e s t i m a t e s of z e r o . sample s i z e s y i e l d of  the data  trapping  rapid  this  for estimating  and  less  placement of  Since v e r y low  i n a c c u r a t e MCP home range e s t i m a t e s r e g a r d l e s s  source,  s u f f i c i e n t data  have  does  home  not  range.  argue a g a i n s t the use of However,  accumulating  t o p r o v i d e a c c u r a t e home range e s t i m a t e s labour-intensive  using  telemetry  Is more  than  using  trapping. Another d i f f i c u l t y t r a p p i n g s t u d i e s of home  t h a t i s f r e q u e n t l y encountered range Is  that grids  c o n t a i n the m a j o r i t y of home ranges  of animals  a r e too  When these  small to  t h a t i n h a b i t them,  so t h a t home range e s t i m a t i o n i s i m p o s s i b l e (Broome 1986).  in grid-  and McMahon,  problems can be a v o i d e d , g r i d - t r a p p i n g i s an  adequate means of home range e s t i m a t i o n i f the r e s e a r c h e r i s o n l y interested home range,  i n the s i z e  of home  ranges.  such as use d i s t r i b u t i o n  i s the s u p e r i o r method.  When other a s p e c t s of  a r e of  Interest,  telemetry  71 LITERATURE  CITED  Aniskowitz, B.T. a n d J . V a i l l a n c o u r t . 1 9 7 9 . A g o n i s t i c t i o n s among w i l d e a s t e r n chipmunks (Tamlas striatus). Z o o l . 57:683-690.  interacCan. J .  B a n k s , E.M., R . J . B r o o k s , and J . S c h n e l l . 1975. A r a d i o t r a c k l n g s t u d y o f home r a n g e and a c t i v i t y o f t h e brown lemming (Lemmus t r i m u c r o n a t u s ) . J . Mamm. 56:888-901. Blair, W.F. 1942. S i z e o f home range and notes history o f t h e woodland deer-mouse and e a s t e r n n o r t h e r n M i c h i g a n . J . Mamm. 23:27-36.  on t h e l i f e chipmunk i n  B l a i r , W.F. 1943. P o p u l a t i o n s o f t h e deer-mouse and a s s o c i a t e d s m a l l mammals i n t h e m e s q u i t e a s s o c i a t i o n o f s o u t h e r n New M e x i c o . Contributions of the Laboratory of Vertebrate Biology, U n i v e r s i t y of M i c h i g a n 21:1-40. B l u e w e i s s , L . , H. F o x , V. Kudzma, D. N a k a s h l m a , R. P e t e r s , a n d S. Sams. 1978. R e l a t i o n s h i p s between body s i z e a n d some l i f e h i s t o r y p a r a m e t e r s . O e c o l o g i a 37:257-272. Boonstra, R. 1 9 8 5 . 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S o c i o b i o l . 3:397-427. Y e r g e r , R.W. 1953. Home range, t e r r i t o r y , and p o p u l a t i o n chipmunk i n c e n t r a l New Y o r k . J . Mamm. 34:448-458.  of the  76  APPENDIX: SOURCE CODE LISTING ( i n BASIC) OF RANDOM CHIPMUNK ENCOUNTER SIMULATION MODEL  TM  DII IA(13,1),B0(13,4),SC|),2) FOl A»5 TO ):F0t l-i TO 2:II1D J C ( A , B ) : H X T : I I I T D H 1 10,1),10,9,20,17,0,21.5,21,),21,21,).5,20.5,20.5 FOl A * l TO 13:F0I 1>0 TO 1:IEAD I A ( A , l ) : I B I T : F O I l < l TO 4:IEA0 BO(A,l):BBXT:l T DAT! 102,1,12,44,13,33: DATA 12,1,11.5,27.5,14,21 DATA ) l , l , 3 , 3 0 . 5 , 0 , 3 0 : DATA H , l , 15.5,31.5,15,32 DATA 00,1,27,25,20,25: DATA 13,1,13,45,15,40 DATA 52,1,1,22.5,1,15: DATA 7,1,2).5,10.5,21,11 DATA (7,1,4.5,25,5,26: DATA 60,1,15.5,51.5,15,50 DATA 10,0,0,30,7,32: DATA 101,0,0,1).5,5,20 DATA 2,0,10,41,13,41 0 C U : IIPUT 'Vbick nontk? *,HO:Iir0T ' l o v B a i y c k l p n n k s ? ",IC 0 DIM t l ( I C , 4 ) , S O ( I C , l ) , I ( I C ) , T ( I C ) , T G 0 ( I C ) 0 FOl 1=1 TO I C : P i l a r i s * c h i p m k  I'»STIMA);:IIP«T I I  0 FOl l«l TO 13:IF IDOIA(B,0) Till 110 0 IF IA(I,1)=1 TIED SS(A,0)=SC{IO,0):SS(A,1)=SC(IIO,1):SI(A,0)=12 0 i r I A ( B , 1 N TIKI SS(A,0)>SC(NO,l}:SS(A,l)*SC(MO,2):SH(A l)=( 0 FOR C=l TO 4 : S I ( l , C ) i I 0 ( l , C ) : I I I T 0 IIIT:IEIT 2 IIPUT ' O i t p t t fr«q«ency? \ n 1 IIPUT 'Output f i l e i a i e ? *,F0 4 OPEI F t FOl OUTPUT IS 11 5 BI=2:IIP0T 'Eocoioter tadl«? (2) ,RI:P8IIT 0 III aov l i l t l a l parameters have beet d i s t i l l e d I n t o : 0 I EM sk: 0 = u i l i a i r a d i o s , l=«est * , 2 * i e s t y,3*centre i , 4 * c e i t r e y 0 I E I s s : l = a c t l v e M a e , 1 - r e t l r e . 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