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Seasonal movements of black-tailed deer on northern Vancouver Island Harestad, Alton Sidney 1979

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SEASONAL ON  MOVEMENTS OF  BLACK—TAILED DEER  NORTHERN VANCOUVER  ISLAND  by  ALTON B.Sc, M-Sc,  SIDNEY 3ARESTAD  University University  of B r i t i s h of B r i t i s h  Columbia, Coluabia,  1971 1973  THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in FACULTY OF GRADUATE ( Faculty  STUDIES  of Forestry  )  He a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o the r e q u i r e d standard  THE  UNIVERSITY OF BRITISH COLUMBIA April,  (c)  Alton  Sidney  1979  Harestad,  1979  In  presenting this  thesis  an a d v a n c e d  degree at  the L i b r a r y  s h a l l make  I  f u r t h e r agree  for  the U n i v e r s i t y it  freely  fulfilment of of  British  available  for  that permission for extensive  the  requirements  Columbia, I agree reference  copying of  and this  this  written  representatives. thesis  for  financial  is understood that gain shall  permission.  Faculty o f  Forestry  The U n i v e r s i t y  of  British  2075 Wesbrook Place Vancouver, Canada V6T 1W5  Date  It  tTuAC  IS '?'?'? 1  Columbia  not  copying or  for  that  study. thesis  s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e Head o f my D e p a r t m e n t  by h i s of  in p a r t i a l  or  publication  be a l l o w e d w i t h o u t my  Abstract Columbian cplumbianus region  on  black-tailed  Richardson)  northern  were m o n i t o r e d  use.  Deer  centres deer,  Resident their  resident  Altitudinal  their  movement  in their  overlapped. from  low  occurred  moving between a s m a l l t r i b u t a r y  main  deer  selection  by  as d e t e r m i n e d of  movements o f b l a c k - t a i l e d deer  deer  Horizontal  ranges moving  migrations  valley  and  the  by  deer  more a v a i l a b l e  energy  and  be  from  habitat  nutrients,  and  lower  c a u s a l d i f f e r e n c e s between v e r t i c a l  m i g r a t i o n s as w e l l a s s e a s o n a l  c e n t r e s can  result  s e a s o n a l l y moving t o more f a v o u r a b l e h a b i t a t s  p r e d a t i o n . The  horizontal  these  home  and  valley. Seasonal  risk  habitats.  by  migratory  spring  winter  occurred  home r a n g e  In  both  between h i g h and by  elevation  migratory  s p r i n g and  migrations  These habitat  shifts  were s e p a r a t e d  although  Columbia.  movements and  or  s e a s o n a l home r a n g e s  home r a n g e s ,  overlapped.  seasonal  hemionug  i n a deep s n o w f a l l  British  made s e a s o n a l  summer home r a n g e s  winter  Island,  either  deer  (Qdocoileus  radio-tagged  to determine  exhibited  but  were  Vancouver  deer  patterns.  deer  r e s o l v e d by  shifts  in  home  a model o f h a b i t a t s e l e c t i o n  and  range  based  on  factors. The.  Amabilis  densities Fir -  of  Twisted  available  S t a l k , and  associations  are comparable with  serai  T h i s abundance o f  stages.  habitats  suggests  not a f f e c t  deer  are  In  met.  that forest  dry  M o u n t a i n Hemlock those deer  i n the food  shrub  in  high  matter  in  Copperbush and  conifer  elevation  h a r v e s t i n g i n high e l e v a t i o n s w i l l  populations, i f their areas  digestible  other habitat  where f o r e s t e d summer r a n g e s  requirements  already  exist,  low  elevation logging  Fern  -  Western  associations closer  to  ranges.  and  Hemlock, provide  their  winter  of  redistribution of  these and  not  Amabilis  Salal  Sword F e r n  will  Use  Logging  of  Western  deer with  Red  sources  ranges than  the  food  sources  may  an  increase  in  of  high  of t h e i r  need  Deer  Plagiothecium abundant  food  e l e v a t i o n summer  result the  Hemlock a s s o c i a t i o n s may  because  Cedar,  Hemlock -  in  deer  F i r - W e s t e r n Hemlock, S a l a l  - Western  deer p o p u l a t i o n s  and  - Western  -  be  f o r these  only  a  population. Douglas-fir,  detrimental habitats  to  during  winter. Management winter  range  policies could  be  emphasizing  detrimental  management f o r b l a c k - t a i l e d winter  r a n g e as  range p r o v i d e s nutrients,  w e l l as deer  i t may  d e e r as i s s e v e r e The  early  t h a t few  snowfalls.  appear unnecessary  severe  to deer p o p u l a t i o n s .  Habitat  winter  range. Because mild  mild winter  g r e a t e r amounts o f a v a i l a b l e e n e r g y  as important  winter  m o b i l i t y of  snow s u g g e s t by  be  of  d e e r must i n c l u d e p r o v i s i o n o f  severe  with  preservation  t o the  over-winter  and  survival  of  sensitivity  to  range.  black-tailed d e e r would be  d e e r and trapped  Corridors joining  to f a c i l i t a t e  deer  their in  high  h i g h and  migrations.  low  elevations elevations  iv  Table of Contents  Abstract  i i  List  Of T a b l e s  . ...  List  Of F i g u r e s  •  ...  Acknowledgement 1.  3.  v i i •  Introduction 1.1.  2.  vi  Scope  Materials  lx 1  And O b j e c t i v e s  2  And Methods  8  2.1.  Study Area  2.2.  Study P e r i o d  11  2.3.  Climatic  12  2.4.  Habitat  2.5.  Effect  2.6.  Tagging  2.7.  Locating  2.8.  Spatial  2.9.  Track Counts  2.10.  Night  2.11.  Sample  8  Data Assessments  12  Of Snow On The P o s t u r e Of S h r u b s  15 16  Tagged  Deer  16  Analysis  18 ,  19  Counts  20  Size  20  Results  22  3.1.  Snowpack I n The S t u d y A r e a  22  3.2.  Abundance  25  O f Deer Food  3.2.1.  Abundance  Of Food D u r i n g  Summer  3.2.2.  Abundance  Of Food D u r i n g W i n t e r  ...............  3.3.  D e n s i t y Of A v a i l a b l e D i g e s t i b l e  3.4.  S e a s o n a l Movements Of K a d i o - t a g g e d Deer  3.4.1.  Statistical  Dry M a t t e r  P a r a m e t e r s Of Home Ranges  40 44 57 60 63  4.  5.  3.4.2.  Home Range S i z e  66  3.4.3.  Position  3.4.4.  O c c u p a n c y Of S e a s o n a l Home Ranges  3.4.5.  T o p o g r a p h i c a l F e a t u r e s Of S e a s o n a l  Of Home Ranges  3.5.  Indirect  Measures Of S e a s o n a l  3.6.  Seasonal  Use Of H a b i t a t s  ,  89 93  Home Ranges 101  Movements ., .  Discussion  108 .120 ...136  4.1.  Size  Of Home Ranges  4.2.  Seasonal  4.3.  S e a s o n a l Movements And S o c i a l O r g a n i z a t i o n  154  4.4.  M o d e l O f S e a s o n a l Movements  156  4.5.  S e a s o n a l H a b i t a t s And F o r e s t H a r v e s t i n g .,  4.6.  Migration Corridors  Literature  Appendix  I  Movements And N u t r i t i o n  Cited  ..136 ...,138  ^..163 168 171 181  vi  List  of Tables  I.  Food A v a i l a b l e  II.  Radio-tagged  III.  Statistical  IV.  S i z e Of S e a s o n a l Home Ranges  67  V.  D i f f e r e n c e s I n S i z e Of Home Ranges  86  VI.  D i s t a n c e s Between  91  VII.  E l e v a t i o n s O f Home Ranges  VIII.  Slope  IX.  Use Of F o r e s t e d And C u t o v e r  X.  D a y t i m e Use Of C o n i f e r In  XI.  I n H a b i t a t s D u r i n g Summer  ..  Deer And S e a s o n s M o n i t o r e d Characteristics  Of S e a s o n a l  61 Home Ranges  S e a s o n a l Home Ranges . . . . . . . . . . . . .  64  102  And A s p e c t Of Home Ranges  . .. 106  Habitats  Serai  123  S t a g e By R e s i d e n t OFL62  S p r i n g And Summer  D e n s i t y Of D i g e s t i b l e D r y  27  132 Matter  Summer On S e a s o n a l Home Ranges  Available  During 148  vii  List  of Figures  1.  Study Area  .  2.  Snowdepths On Mount C a i n  23  3.  Percent  32  4.  A n n u a l Growth And Stem H e i g h t  5.  Abundance Of Food  Available  I n Summer  41  6.  Crown D e p t h And Stem H e i g h t  Of S h r u b s  45  7.  Vertical  8.  Food  9.  Density  10.  Seasonal  Home Ranges Of A l t i t u d i n a l  M i g r a t o r OFL61 .. 70  11.  Seasonal  Home Ranges Of A l t i t u d i n a l  M i g r a t o r OFL68 .  72  12.  Seasonal  Home Ranges Of A l t i t u d i n a l  M i g r a t o r OFL67 .  74  13.  Seasonal  Home Ranges Of A l t i t u d i n a l  M i g r a t o r OFL60 .  76  14.  Seasonal  Home Ranges Of H o r i z o n t a l M i g r a t o r OFL71  15.  Seasonal  Home Ranges Of R e s i d e n t  OFL58 . . . . . . . . . . . . .  80  16.  Seasonal  Home Ranges Of R e s i d e n t  OFL62 . ,  82  17.  Standard  Distance  18.  P e r i o d s Of O c c u p a n c y Of S e a s o n a l  19.  Winter  Cover  ,  9  And Biomass Of L i c h e n  Distribution  Available  Of S h r u b s  Of Shrub  35  A n n u a l Growth  In H a b i t a t s D u r i n g  47  Winter  53  Of D i g e s t i b l e D r y M a t t e r  D e v i a t i o n And S i z e Of Home Range  M i g r a t i o n s Of OFL68  Snowfalls  58  ..  .  Home Ranges  78  84 94  And 0FL61 I n R e s p o n s e To  ...  98  20.  Seasonal  Movement P a t t e r n s I n S t u d y  21.  E l e v a t i o n Of H i g h e s t  22.  Night  C o u n t s I n The Mount C a i n S e c t o r . . . . . . . . . . . . . . 114  23.  Night  Counts  24.  Use Of F o r e s t e d And C u t o v e r  25.  Habitat  Deer T r a c k  Each  Habitats  Season  109  On Mount C a i n Road . 1 1 1  I n T h e Croman S e c t o r  Use D u r i n g  Area  118 121 126  viii  26.  H a b i t a t Use D u r i n g  Summer By  Deer 27.  Model Of S e a s o n a l  Migratory  And  Resident  o .......... Movements  129 157  ix  acknowledgement Dr. F . L . B u n n e l l p r o v o k e d this  thesis  motivation. Kimmins,  and  I  am  Thanks a l s o  J.H.  Myers,  many o f t h e c o n c e p t s d e v e l o p e d i n  indebted  to  him f o r h i s g u i d a n c e and  go t o D r s . H . D . . F i s h e r , and D.M.  K. Graham,  J.P.  S h a c k l e t o n f o r t h e i r a d v i c e and  direction. Sincere Division  of Canadian  Chester,  From t h e  objectivity  and  and  collection  Richard  S m i t h , George T a y l o r ,  their  Jim  i s appreciated  project  made  Hans  Ian  research.  Stan  K a p i t a n y . The and  I  thank  and e n c o u r a g e m e n t . Smith  through  Anthea  Farr  his t o my  by  her  to the design of the project  Dr.  Don  D a r y l l Hebert, and  Logging  substantial contributions  contributed  o f d a t a . I thank Dr.  Englewood  hospitality  in wildlife  ideas  Ellis,  discussions.  this  insight  and a t t i t u d e  enthusiasm and  of  the  J o n e s , and J u l i u s  o f Woss f o r t h e i r  inception  to  P r o d u c t s Co. L t d . , p a r t i c u l a r l y  g i v e n by Danny J o h n s o n  residents  approach  Forest  extended  G o r d o n F l o w e r d e w , Greg  inspiration the  gratitude i s  Eastman,  Byron  Weber  R o c h e l l e and Susan  Allan  Edie,  Mason, Bud P r a t t ,  for  their  Stevenson  Bud  enlightening  p e r m i t t e d use o f  data. Canadian  Fellowships, B.C. F i s h  and  Forest  P r o d u c t s Co. L t d . , H.R.  the W i l d l i f e Wildlife  B.C. p r o v i d e d f i n a n c i a l  Research Branch,  and l o g i s t i c  Division and  MacMillan and R e g i o n  the  support.  Family 1 of the  University  of  1  1.  Introduction The  Columbian  columbianus  will  the c o a s t a l  deer  deer,  be r e f e r r e d  t o as b l a c k - t a i l e d  r e g i o n o f North  54°N  in  British  America  Columbia  ( 0_. h ^ s i t k e n s i s Merriam  accessibility  of b l a c k - t a i l e d  from  Early  deer  forests  provided l i t t l e allowed  forest  snow c o v e r  was  luxuriant  growth o f  1968,  Smith  where snow d e p t h s  ephemeral.  The  1968).  deer In  in  authors  were  s h a l l o w and that  deer  extensive  populations.  h a d no r e a s o n t o winter  habitat  C o a s t a l Western Hemlock a n d t h e s u b a l p i n e  Mountain  i n areas  Edie  they  1950,  studies  recognized  would be r e q u i r e d a s  o f deep s n o w f a l l .  Hemlock b i o g e o c l i m a t i c z o n e s Island,  Leopold  deer  and  beneficial  c o v e r , and t h a t  climates,  of the  herbs,  were r e l a t i v e l y  f o rsecurity  milder  mature f o r e s t s  the  removal  These  c o u l d be d e t r i m e n t a l t o b l a c k - t a i l e d  that  most  understory  t o be h i g h l y  logging  for  vegetation  working  shrub  grasses,  tall  expect  the  deer i n h a b i t e d  Since  needed  However,  of  (Cowan 1945, E i n a r s e n 1946,  Gates  i n areas  and  food f o r deer.  h a r v e s t i n g was c o n c l u d e d  species  1961,  conducted  i t one  o b s e r v a t i o n s were made i n t h e C o a s t a l D o u g l a s - f i r  canopy  Brown  the S i t k a  (Cowan 1945, E i n a r s e n 1946, Brown  w h i c h have minor amounts o f h e r b  t h i s deer  to  Columbia.  forests  to  northern C a l i f o r n i a  makes  These  hence,  It inhabits  ) (Cowan 1956).. The abundance and  1961).  shrubs,  deer.  s t u d i e s r e p o r t e d t h a t few b l a c k - t a i l e d  the c o a s t a l old-growth  forest  hemionus  where i t merges w i t h  p o p u l a r b i g game s p e c i e s i n B r i t i s h  and  Odocoileus  ( R i c h a r d s o n ) , i s a s m a l l s u b s p e c i e s o f mule d e e r and  henceforth  about  black-tailed  and  Harestad  ( sensu  Krajina  1965) on  (1971) and Smith  Vancouver  and D a v i e s  (1973)  2  observed  that  black-tailed  forests  far  from  snowfall, deer  (Robinson 1972,  1956,  1978).  sought Bloom  after  and  by  the  forest  deer  populations  Their r e s u l t s could  indicate  disparities  forest  understood.  created  1973,  Jones  easy  on  deer  between be  effects  concerning  Scope And  Bunnell.  deer,  and  1975,  1974  and  (1974,  harvesting  thus  1975) on  and  blackcover.  winter  ranges  deer  of  that  are  forest  not  attributed  on  forests  deer  to a lack  movements, f o o d r e q u i r e m e n t s , deer  clearly  harvesting  of old-growth and  effects  has  resources. of  basic  habitat  use,  .  Objectives  In response tailed  Smith  f r e q u e n t l y old-growth  u s e r s of timber  partially  1965,  abundance.  black-tailed  p o p u l a t i o n dynamics of b l a c k - t a i l e d  1.1.  Guiguet  locally  w i t h deep w i n t e r snow  c o m p e t i t i o n f o r use  These c o n f l i c t s c a n  survival  among t h e s e s t u d i e s i n d i c a t e on  deeper  black-tailed  accessibility,  of f o r e s t  i n areas  Controversy over  conflicts  knowledge  and  by  that h a r v e s t i n g of f o r e s t e d  harvesting  abundance and  Cowan  are  pristine  r e g i o n s of  i n d u s t r y . Jones  have a d e t r i m e n t a l e f f e c t The  some  winter range  ranges  wood volume and  these  to the deer's  Game  winter  In  as  1956,  (1978) c o n s i d e r e d e f f e c t s  tailed  and  critical  Edwards  These  f o r e s t s of high  deer  t o be  were a b u n d a n t i n  areas.  h a b i t a t s used  A l a s k a Dept. F i s h  Bloom  of  logged  forested  were t h o u g h t  deer  a  t o t h e need f o r more i n f o r m a t i o n series  of p r o j e c t s  Movements of b l a c k - t a i l e d  thesis  while  complementary  habits,  nutritive  quality  deer  studies  of the  was  about  initiated  are are  f o r a g e , and  black-  by  Dr.  described  in  this  with  food  abundance  and  concerned the  F.  L.  3  distribution No  of lichens  p u b l i s h e d s t u d i e s have been made on t h e s p a t i o - t e m p o r a l  distributions forest  of  free  ecosystems.  have been  on E o c k y  but  these  habitats  i n constrast  Columbia  occupies  temporal  ranging  has  studies  forested  been  structure  Mountain  (Estes  mammalian  social  increasingly mammalian Geist  focused  1974, Jarman  behaviour  1974, Crook Habitat  in  British  on for  forest  in  because  requirements  have  extrapolation  from d e e r  examine  the  consider  their  spatioof upon  Understanding  systems  theoretical  which  have  of  become  considerations  of  1970, A l e x a n d e r 1974,  deer  i s of p a r t i c u l a r  practises,  been  of b l a c k - t a i l e d  of  studied,  some  populations i n differences  Columbia dispersion  in  deer  of  the  is  deer's  necessary  aspects of habitat  have  been  southern climate  and and  and t h e s e r e g i o n s , i t i s patterns  interest  and b e c a u s e i n f o r m a t i o n  management p l a n s . A l t h o u g h c e r t a i n  between B r i t i s h  in  dependent  of the modifications t o  made by f o r e s t r y  Because  British  organization,  be h i g h l y  ( E i s e n b e r g 1966, C r o o k  requirements  regions.  in  are important t o the understanding  use by b l a c k - t a i l e d  the habitat  Variation  i n open  e t a l . 1976).  Columbia  environment  coastal  living  which  1974).  organizational upon  deer  social  to  in  0^. h.. h e m i o n u s  o f deer  habitats.  particularly  spatio-temporal d i s t r i b u t i o n s of  mule d e e r ,  were  demonstrated  deer  and s e a s o n a l movements  to the b l a c k - t a i l e d  distribution,  ungulates  black-tailed  S t u d i e s o f home r a n g e  conducted  (Eafinesgue),  habitat  and s h r u b s .  assumed by snow-free vegetation  necessary  black-tailed  s e a s o n a l movments o v e r t h e whole y e a r .  deer  to and  4  Seton over  (1909) w r o t e t h a t  t h e c o u n t r y ; each  actual  animal  i s  refugia,  fitness  at  localization  of  an  o f an a r e a . T h i s f a m i l i a r i t y  knowledge  of  roams  of  food  animal"s  i t s efficiency  evading  would  of finding  p r e d a t o r s and u l t i m a t e l y 1953,  f o o d and i t s increasing i t s  Metzgar  1967,  Burt  by  ( 1 9 4 3 ) , I c o n s i d e r home r a n g e  an a n i m a l d u r i n g i t s n o r m a l  with feeding, r e s t i n g , Seasonal  t o be t h e a r e a  activities  r e p r o d u c t i o n , and a v o i d a n c e  movements i n c l u d e  both  associated  of predators.  m i g r a t i o n and s e a s o n a l s h i f t s i n  t h e c e n t r e s o f home r a n g e s . M i g r a t i o n i s t h e p e r i o d i c from,  and  moves f r o m ranges them  return  t o , an a r e a . T y p i c a l l y ,  one s e a s o n a l home r a n g e  o f a m i g r a t o r y a n i m a l aire s e p a r a t e d , are  used  mainly  f o rtravel.  time  shifts  that  home  intensity  range  d i f f e r from  i s  of use of  used  The' s e a s o n a l home  and  relative  zones  of  between  t o the  a t each  migrations i n that  throughout  portions  animal  The a n i m a l s p e n d s l i t t l e  i t s p e n d s i n t h e home r a n g e s  i n home r a n g e  departure  the migrating  t o another.  t h e z o n e between s e a s o n a l home r a n g e s  same  Covich  C r o o k e t a l . . 1976).  traversed  of  give  s o u r c e s , p r o t e c t i v e c o v e r , and  ( D a v i s e t a l ^ 1948, B l a i r  Following  in  random  t o have a d a p t i v e v a l u e by p r o v i d i n g t h e  thus i n c r e a s i n g  capability  1976,  proposed  with f a m i l i a r i t y  animal  animal  h a s a home r e g i o n , e v e n i f i t h a s n o t an  home". T h i s phenomenon o f  activities  the  "no w i l d  the  the home  time  amount  end. S e a s o n a l much o f t h e  y e a r . However, t h e range  i s  altered  of activities i s beneficial  because  seasonally. Although of  localization  the f a m i l i a r i t y  animal's  gained  requirements  can  of  the  home  range,  not  a l l  the  be o b t a i n e d a t t h e same p l a c e . T h i s  5  will  be  especially  availability over  an  of r e s o u r c e s . Animals  area  to s a t i s f y  movements, and related  to  true i f there are  hence t h e s i z e  their  energy  relationship  occurring  productivity  (McNab  Harest'ad and expands  their  increases  Since the s i z e  requirements location  o f home r a n g e  of  in  trophic  and  productivity  Turner  may  these  also  be  the  habitat  range  size  and c o n t r a c t s  ( H a r e s t a d and on  df energy,  is  e t a l . . 1969,  home  depends  availability  o f t h e home r a n g e  and  requirements,  of home' r a n g e  the  extent of  status  1968,  energy  t o move  with m o d i f i c a t i o n s t o  Schoener  in  the  vertebrates,  B u n n e l l , i n p r e s s ) . F o r mammals,  with  in  are t h e r e f o r e obliged  requirements  1963,  changes  r e q u i r e m e n t s . The  because  with i n c r e a s e s i n h a b i t a t press).  temporal  Bunnell, i n  the  animal's  i t follows  influenced  by  that these  factors. The has  link  been p r o p o s e d  More  f o r o v e r 80  recent studies  Bertram  and  statement  sources  Bergerud  Rempel  1977)  have  not  this  causal  (Byford  1969,  potential  1972),  but  no  Some  studies  have  differences  between  the  ( D e a l y 1959,  Klein  seasonal  1965  the has  movements. I n a r e a s  year melted,  or  during  quality  beyond  the  studies  have  local  food  in  and  1970,  main p r o c e s s e s c o n c e r n i n g t h e f o o d s u p p l y  throughout snowpack  made.  nutritional  seasonal ranges  explain  was  1974,  Other  changes  1919).  LeResche  progressed  with  Strandgaard  Adams  1974,  relationship.  movements o f u n g u l a t e s  s e a s o n a l movements  (Morgan 1896,  1970,  the food supply  Two  years  and  (Mackie  of  described  of  between f o o d a v a i l a b i l i t y  quantification demonstrated f o r a g e s on Hebert are  the  1973). used  to  where f o r a g e i s a v a i l a b l e  spring  and  summer  when  of the forage i s suggested  as  the the  6  most i m p o r t a n t f a c t o r 1965,  Klein  1965  differences  in  inducing  and  1970,  to differences i n  forage  (Cook  1972,  topography,  regions,  following Dixon  1965,  second  American Snow  Longhurst  mobility,  food  and  animals seek  sources,  black-tailed snowfall  deer  regions  r e g i o n s . The  of  pattern  implies  population.  the  1956)  forage  of  Hickey  W i n t e r and  with  period  animal 1932, et a l .  migration i n  North  availability.  foraging  1977  animal's migration,  and  density,  , Telfer  1978).  o f s e a s o n a l movements by populations i n deep  in  low  snowfall  as p r o g r e s s i v e  movements by  high  phenological  Klein  quality  1965) . T h i s  s e r i e s of short and  and  Dalke  populations  ( D e a l y 1959,  would  term  proposed  home  involve  ranges  the  whole  summer home r a n g e s have been o b s e r v e d f o r  population  b u t no  the  (Russell  depth  resident  migratory  a transitional  migratory  a non-migratory Taber  and  by  1960,  the  snow  pattern  moving i n phase  stages  the  and  m i g r a t i o n s are proposed  the p o p u l a t i o n  during  (Leege  that  to elevation  of l o c o m o t i o n . Through  accepted is  mountainous  o f snow on f o o d  out areas of a p p r o p r i a t e  currently  In  1976).  restricts  increases costs  the  differences  of the forage Knowlton  of  due  from  migration  Stelfox  i s the e f f e c t  where f o o d i s a v a i l a b l e The  are related  main p r o c e s s u s e d t o e x p l a i n  ungulates  covers  1973,  nutritive  condition  can r e s u l t  e t a l . 1952,  Hebert  These  precipitation.  state  (Dalke e t a l .  on s e a s o n a l r a n g e s a r e  altitudinal  t h e most n u t r i t i o u s  Klein The  or  and  differences  induce  1973).  phenological  1965),  temperature,  to  1934,  1965,  the  Klein  phenological  are thought  Hebert  the f o r a g e a v a i l a b l e  largely  in  migratory behaviour  of  black-tailed  deer  (Dasmann  s t u d i e s have been made o f t h e c o m p l e t e  and  annual  7  behaviour  of  migrations  have not  inferred In tailed  migratory been  study,  deer occupying  seasonal  described  Furthermore the a c t u a l  for individual  deer  my o b j e c t i v e s were t o d e t e r m i n e coastal  p r a c t i s e s , do have home  which p r o m o t e Finally  population.  but  from changes i n p o p u l a t i o n d i s p e r s i o n . this  logging  that  a  range  seasonal  use  exists,  ecosystems,  home r a n g e s ,  be  movements  are considered  implemented  management p o l i c i e s  seasonal  in  forestry  t o enhance d e e r  with  modified  some f a c t o r s home  regard  practises  populations.  by  what p a t t e r n o f  and t o e x p l o r e  movements by d e e r between  these  could  forest  i f black-  ranges.  t o changes and  deer  8  2.  M a t e r i a l s And Methods  2. 1.  Study The  km  study  east  area  i s located  o f Woss, B r i t i s h  126°25'W. Biver  Area  The a r e a ,  on n o r t h e r n  Columbia  187 km  is  i n extent,  2  originates  in  the Davie R i v e r  Valley,  the  a steep  topography  sidehill  V a l l e y i s wider than on  both  on t h e e a s t  adjacent  Mount C a i n  rises  side  at  The  Davie  Tsitika area.  Croman  Creek  o f t h e Davie R i v e r i s  o f Croman C r e e k . T h e r e i s  valley  floor  that  e l e v a t i o n , the slope  t o 1000 m b e f o r e  50°12'N  s i d e of the the  20  steepening  rises  to  i s gentle  near the peaks o f  and M a q u i l l a Peak.  Access to the through  sides  to the  between 600 and 700 m. Above t h i s the land  ( F i g . 1).  and d r a i n s t h e n o r t h e r n  Although  s i m i l a r t o t h a t found  centred  i n c l u d e s t h e upper  V a l l e y and t h e Croman C r e e k V a l l e y  River  as  and  Vancouver I s l a n d ,  a  system  study  area  of logging  i s  from  the  roads running  Nimpkish  the l e n g t h  Valley  o f Croman  C r e e k and D a v i e R i v e r . T r i b u t a r y r o a d s l e a d up t h e s l o p e s  of the  m o u n t a i n s t o e l e v a t i o n s o f a b o u t 600 m i n  Croman  Creek  Valley  and  topographical  sectors  are  as h i g h  a s 1150 m on Mount C a i n .  delineated  tagged Croman  deer,  i n Figure and  habitat  (1) and Mount C a i n  represented  in  1. Most o f  the  and  Hemlock  the Alpine  Krajina  the  tagging,  (2) s e c t o r s . The b i o g e o c l i m a t i c  zones  area  were  of  i n the  study  assessments  location  conducted  Zone i n t h e l o w e r e l e v a t i o n s Mountain  Four  are the C o a s t a l  (200  to  900  m) ,  Zone i n t h e u p p e r e l e v a t i o n s  Zone n e a r t h e m o u n t a i n t o p s  Western Hemlock the  subalpine  (800 t o 1600  (above  1 9 6 5 ) . I n Croman V a l l e y where t h e c l i m a t e  1500 m) i s cooler  m),  (sensu and  9  Figure are  Study  unlogged.  dashed 3.  1.  lines  a r e a . Shaded a r e a s a r e l o g g e d ,  S e c t o r s a r e numbered and cr  waterways  M a g u i l l a , 4 . Hoomak.  :  their  1. Croman,  unshaded  boundaries 2.  Mount  areas  shown  by  Cain,  10  11  snowfall closer  greater,  to the  the  s u b a l p i n e M o u n t a i n Hemlock Zone  v a l l e y .bottom t h a n i t d o e s i n t h e  Mount  extends  Cain  (2)  along  the  sector. F o r e s t h a r v e s t i n g began i n t h e flat  valley  forests and  bottom near Davie  were  logged  higher elevation  below  800  above 800 the  m were l o g g e d .  i s the  forest  of  800  The  while  blocks  less  usually  slashburned.  the study  2.2.  than  area  Study  of  81  t o 27  lower  harvesting  was  years,  in  of  the  been c o n d u c t e d . study  the  forests  predominantly  patch  cutting  of  Logged a r e a s  immature  serai  were  stages  in  years o l d .  began i n J u n e  1974  difficulties  attempts  at  tagging  and  with and  continued the  locating  spring  intensively the deer  until was  and the  early  summer o f  spring  reduced  of  1976.  1975. The  i n t h e summer o f  s e a s o n a l h a b i t a t s were made.  The  deer  through  to  May  radio-transmitters  by  of  64%  elevation  r a d i o - t r a n s m i t t e r s were o b t a i n e d and  of  forests  i s logged. In comparison,  recent  the  forests  1 differ  Here,  Reliable the  of  have b e e n c u t . Mount C a i n  of the  ha has  up  39%  w h i l e o n l y 7% o f  sector.  forest  During  1975,  sidehill  Period  Technical  subverted  o n l y 27%  i n more  ranged  Field-work 1977.  By  elevation  of the  f o u r s e c t o r s of F i g u r e  m in elevation  pattern  clear-cutting,  extraction  the f o r e s t s t h a t  (1) s e c t o r has  logged.  then  most e x t e n s i v e l y l o g g e d  below  Croman  and  were l o g g e d ,  The  a r e a i n 1948  R i v e r . Much o f t h e l o w e r  f o r e s t s began.  m in elevation  proportions  (2)  first  study  The  deer  attached deer  intensity 1976  in to  were of  1974. deer  located location  while e v a l u a t i o n s  were  located  until  12  December  2.3.  1976 and t h e p r o j e c t t e r m i n a t e d  Climatic Five  Data  weather  stations  Branch  (B.C. M i n i s t r y  study  a r e a . Monthly  and  continuous  a  recording  was  sponsored  of Environment)  by t h e R e s o u r c e A n a l y s i s were  maintained  minimum a n d maximum t e m p e r a t u r e s  record  thermographs.  precipitation  i n May 1977.  of  temperature  During  recorded,  was  periods  while  with  during  i n the  were  obtained  taken using  no snow, m o n t h l y  periods  with  snow  m o n t h l y snowpack d e p t h s were measured. In  addition  stations,  snow d e p t h  estimated elevation depths  to  every from  were  the  snow  d e p t h s measured  and p e r c e n t o f ground c o v e r e d  two  weeks  at stations  300 t o 1200 m a l o n g sometimes  measured  the  Habitat The  to  and then  Seasonal  be  referred  associations stages if of  necessary,  Cain  road.  Snow during  serai  stages.  transferred  were o u t l i n e d  on t h e  The mature s e r a i  stages  t o as f o r e s t e d p l a n t a s s o c i a t i o n s or, simply,  home r a n g e s  and  s e a s o n a l home r a n g e s .  serai At t h e s e  stages  referred  t o as  serai  were i n s p e c t e d o n t h e g r o u n d a n d ,  the s t r a t i f i c a t i o n  associations  were  a c c o r d i n g t o m a j o r mature p l a n t  a n d t h e immature s e r a i  . Seasonal  deer  home r a n g e s  stratified  a s s o c i a t i o n s and immature will  Mount  100 m o f  Assessments  photographs.  photographs  l o c a t e d every  were  or frequent s n o w f a l l .  l o c a t i o n s f o r each r a d i o - t a g g e d  aerial  by snow  more o f t e n , e s p e c i a l l y  periods o f intense r a d i o monitoring  2.4.  a t t h e weather  was a l t e r e d .  stages sites,  were  Typical  selected  Habitat  examples within the  Assessments  were  13  conducted plant  which  included  s p e c i e s and  estimates  of  their  shrub  a physiographic description,  percent  cover  and  abundance.  Lists  of  estimates of percent cover used  by O r l o c i Plant  (1964) and  their  through  percent cover  Bell  (1971).  correspond classified their  into  percent  classification  hierarchy year the P.  of B e l l  and  aquilinum  percent  of  plant  of  the  associations  and  used  are used  by  shrubs,  herbs.  - recently  The  deer  were  were  (  of herbs  fern  -  bracken  in  less  c o n i f e r s l e s s than  20%  10  used  ascending than  one  i s greater  than  percent  cover  percent cover of herbs  i s g r e a t e r than  by  fern  criteria  l o g g e d and  - percent cover  the  those o f  and  stages  ;  and  that are c h a r a c t e r i z e d  Kuhn), and serai  species  Brooke e t a l . ( 1 9 7 0 ) ,  Cutovers  stages  o f P_. a q u i l i n u m  of  of l i s t s  deer  ; conifer  of  ; shrub -  total  m high i s greater  than  . Forested  two in  (L.)  cover of shrubs  and  methods  radio-tagged  (1964),  (1971).  i s g r e a t e r than  cover  releve  by  h e i g h t of c o n i f e r s ,  : newly l o g g e d  percent cover  the  and  species  used  of these  serai  o l d cutover; herb  - percent  20%  )  forests  names  ( Pteridium aquilinum in  in  Orloci  five  cover  plant  at H a b i t a t Assessment s i t e s t o  Common  to those  distribution,  were o b t a i n e d by  comparison  a s s o c i a t i o n s d e f i n e d by  of  B r o o k e e t a l . (1970) .  associations  were d e t e r m i n e d  list  groups: the  plant those  subalpine  biogeoclimatic their (after  typical Bell  associations  were d i v i d e d  into  i n t h e C o a s t a l W e s t e r n Hemlock Zone and  those  Mountain  zone, the position  1971).  Serai  used  stages  deer  Hemlock  associations on t h e  by  Zone.  Within  were a r r a n g e d  s l o p e and were  their  arranged  each  according to  moisture in  regime  order  of  14  increasing The  s u c c e s s i o n a l age. abundance  sarmentosa  (Ach.)  Assessment s i t e positives that  the  into  of  square  by  one  was  Ach. low  cm  3 = 76-100%. The in  sites,  a relationship  and  independent  was  (1978).  percentage  of  this  abundance  (kg  was Howell  site.  The  proportion  these  classes  of  representative  measured. heights  and  area  the  percentage  for  contributed  by  not these  gridded  cover  i n each  estimate  each  Habitat  measured.  the  For  some  percent  cover  biomass the  26-75%  total  site.  between t h i s  the  of  obtained  by  percent  cover  Assessment  site  other Vaceinium  30 cm  to  v a l u e s as  (including  by i d e n t i f y i n g d e n s i t y at the a t the  site  was  then  density  shrubs  area  f o r Vaceinium  o f each c l a s s  than  an  so  ).  area  shrub  and  lichen  h e i g h t o f e a c h stem  Stems l e s s were  - 1  Colour  a screen  estimated  shrub of  Habitat  1 = 6-25%, 2 =  relationship,  ha  G a u l t h e r i a s h a l l o n Pursh.  four c l a s s e s of r e l a t i v e  an  of l i c h e n  0 = 0-5%,  determined  Using  each  c l a s s e s corresponded  obtained  estimate  t o biomass  Xs. a l a s k a e n s e  selected  The  of t h e c l a s s  was  Alectoria  photographs.  image c o v e r e d  ranges;  was  of  were p r o j e c t e d o n t o  f o r a r b o r e a l l i c h e n s at each  Shrub  and  of l i c h e n  converted  Smith,  The  range  at  oblique a e r i a l  c l a s s e s were a s s i g n e d  the  mainly  estimated  0 t o 3.  cover  cover  estimate  was  canopy  squares.  percent  Stevenson  consisting  p o r t i o n of the  percent  mid-point  ,  level,  c l a s s e d from  following  an  lichen,  the f o r e s t  central  450  of  of shrub rooted  i n height Estimates  were made by  paryifolium  between Habitat  one  and  Assessment  exhibiting  each  of  In  a  estimated.  density, a plot in  the  were c o u n t e d of  species) ,  the  clipping,  plot but  was was  their  amount o f  food  drying,  and  15  weighing  the  heights.  Species-specific  growth  growth  relationships  contributed  2.5.  annual  of  shoots  regressions  were  then  used  cover  availability  depends  accumulation newly  on  of shrubs  of  the  to  estimate  effects  were t o be i n v e s t i g a t e d ,  mild  little  snow  subsequent by  ascent  Some  except  insight  of shrubs  change  in  height  of  J u n e 6, 1976, an a r e a and  a shallow  snowpack  m southeast  hole  was  second  was c h o s e n  that  the was  measured.  bifurcation, shrub. the  tape  Some  stems  from  winter.  2, 1976, a f t e r  stems r e g a i n e d t h e i r  o f 1976 the  stem  shrubs area  A small  was e n c o u n t e r e d .  was a t t a c h e d t o t h e stem  and  directly  d i s t a n c e between t h e marked  On  i n the f o r e s t  weather s t a t i o n .  a shrub  two  A a  below  markers  above t h e f i r s t  t h e snow had c o m p l e t e l y  snow-free  and  The  w h i l e o t h e r stems were marked i n t h e crown  On J u l y  i n the  measuring  Stalk association  were  very  depression  was s e c u r e l y p l a c e d on t h e g r o u n d  marked stem. The v e r t i c a l  winter  had an abundance o f  o f t h e 900 m Mount C a i n  of  t h e snow had m e l t e d .  (30 cm) r e m a i n i n g  dug i n t h e snow u n t i l  marker  the  t h e marker a f t e r  numbered p i e c e o f f l a g g i n g  1976-1977, was  t h e snow and t h e n  was i n an A m a b i l i s F i r - T w i s t e d 250  effect  was o b t a i n e d i n t h e s p r i n g  marking stems b u r i e d beneath  snow  s t e m s . The w i n t e r  during l a t e  into  of  stems t o snow  I was n o t a b l e t o measure t h e  snow on t h e d e p r e s s i o n o f s h r u b  elevations.  food  Assessments.  r e a c t i o n s of shrub  fell  the  during periods  when t h e s e  higher  height:annual  Of S h r u b s  to deer  structural  and melt.  fallen  and  stems o f v a r i o u s  by a l l stems measured i n t h e H a b i t a t  E f f e c t Of Snow On The P o s t u r e The  from  of  major the  m e l t e d and  p o s t u r e s , h e i g h t s between t h e  16  markers  were  Vaceinium  alaskaense  Smith  were a l s o  2.6.  Tagging Deer  ensure were  remeasured.  the  shrubs  several  from  different  movements under  different  Most  were  Menziesia.  of  these  immobilized  environmental  using  a  Darts  Chemical  Co.)  D a r t I n c ) . The i n single  gate C l o v e r t r a p s  Radio-transmitters to  selected  eartagged attached,  deer  (Wildlife  a d u l t s and y e a r l i n g s . with  a plastic  flap  that  when  relaxant,  form  o r i n powdered f o r m  remaining  conditions  muscle  administered i n l i q u i d  captured  f erruginea  d e e r were f r e e - r a n g i n g  chloride,  (Pneu  were  p a r t s o f the study a r e a to  succinylcholine (Palmer  marked  marked.  represented. and  of  although  were t a g g e d  that  tagged  ,  Most  by  Capture  by Pneu D a r t s  were: t a g g e d , ,  (Clover  were  1956) .  M a t e r i a l s I n c . ) were a t t a c h e d The  radio-tagged  on which  and w i t h numbered R o t o t a g s  deer  a reflective  (Nasco Co.)  or  were  number metal  was clip  tags.  2.7.  L o c a t i n g T a g g e d Deer Radio-tagged  depending the  were l o c a t e d  deer  periods, being  s e s s i o n i n August hours  in  weekly  located  on  various  o f t h e d e e r and t y p e and  o b t a i n e d from  at least  During c e r t a i n  two  were  upon a c c e s s i b i l i t y  information  selected  deer  the l o c a t i o n s .  when  I  was  in  schedules value  Radio-tagged the  study  l o c a t i o n s were made more f r e q u e n t l y located  every  to  determine  deer area. with  12 h o u r s . D u r i n g a t e n day  1975, a r a d i o - t a g g e d d e e r was r e l o c a t e d  order  of  the  optimum  every  schedule f o r  17  monitoring  movements.  Radio-tagged  deer  were l o c a t e d u s i n g a whip a n t e n n a  hand h e l d , t h r e e e l e m e n t for  direction  of  The Y a g i the  r a d i o - t a g g e d deer time,  my l o c a t i o n ,  When  tagged  time,  location,  deer  Bearings obtained deers'  the  resulted  depending  from  from  as upon  This accuracy vertices  different  of  a  signal with  being the index  the  and  date,  comments.  t h e t a g number,  than  1975, r e l i a b i l i t y from  their  and confirmed  were  date,  quickly  The r a d i o - t a g g e d  triangulation.  Bearings  were n o t i n c l u d e d  Aberrant  study  of  bearings  a r e a . The l o c a t i o n "rough",  intersection  triangle  m  (very  class.  in  the  The  "very  of the  During  two  were w i t h i n a  rough).  o f the telemetry  locations.  or  100 m ( a c c u r a t e ) , 100 t o  250  deer  detected  on whether a t l e a s t  intersection  within the accurate  signals  by  was b a s e d  fell  visited  signal,  locations.  degree  estimates  triangulating  When  the t a g number,  "accurate",  greater  taking  a  known l o c a t i o n s .  direction  the  and  by  from  the  b o u n c e s and were e a s i l y  of a r a d i u s o f l e s s than  of  used  determine  location.  (rough),  summer  to  the other bearings  of  familiar  rough"  signal  estimated  estimate  was j u d g e d  circle  radio  were  estimate  the  from  were s i g h t e d , I r e c o r d e d  of  became  bearings.  deer  used  deer  a  and comments.  final  I  was  b e a r i n g of t h e r a d i o  different  frequently  of  radio-tagged  of radio-tagged  were l o c a t e d , I r e c o r d e d  locations  the  after  antenna  a t three t o f i v e  distinctly in  Y a g i a n t e n n a . The whip a n t e n n a was  p r e l i m i n a r y s e a r c h and l o c a t i o n  moving v e h i c l e .  and  250  m  Most o f t h e  the spring  and  system  was t e s t e d  and  immediately  field  estimated  by o b s e r v a t i o n o f t h e t a r g e t  locations deer.  were After  18  many  locations  triangulation  were  confirmed,  I  was  methods were g i v i n g good  satisfied  estimates  that  of  a  the  deer's  locations. In  the  coordinates which on  field,  was used  bearings  performed.  using  (3.16 cm t o 1.0 km) w i t h  t o l o c a t e tagged deer  were t a k e n .  a 2.08 cm t o  locations  a map  100  m  Estimated o f tagged  a series  Bearing  forest  deer  were  of origins.  map  of  recorded  origins  were s t a n d a r d i z e d  the  polar coordinates  use  i n plotting  2.8.  Spatial Home  there  are  an  100%  an  be  daily,  animal  home  closest  90%  Consideration  to a single  to rectangular  a  specific  seasonal,  polar  deer  and  coordinates  so that  e a c h was  error.  These  common o r i g i n and coordinates  for  data.  or of  of  only  a  the  animal's  interval.  Thus,  home r a n g e s .  Home  locations  t i m e p e r i o d . The a b s o l u t e o r  a l l the  expressions 50%  o f an  as a p r o p o r t i o n o f t h e known  includes  Other  time  and l i f e t i m e  within a particular  range  individual.  as  integrated expression  over  r a n g e s c a n be c o n s i d e r e d for  located  Analysis  and movements  can  radio-tagged  minimizing  and a n a l y s i s o f l o c a t i o n  ranges  locations  converted  from  and t r i a n g u l a t i o n s  O r i g i n s were p l a c e d  near a c o n c e n t r a t i o n o f l o c a t i o n s , thus "local"  locations  p o s i t i o n s were t h e n  cover  locations  and  rectangular  known  locations  of  an  o f home r a n g e s i n c l u d e o n l y t h e individual's  known  p o r t i o n o f the observed  useful i n identifying  cores  of i n t e n s e  unusual  movements  whose  inclusion  estimates  of home r a n g e  parameters.  locations. locationsi s  u s e , and f o r e l i m i n a t i n g would  create  erroneous  19  L o c a t i o n s and movements as  home  ranges  statistical elegant,  and  methods  do  not  information.  On  c a n be p r e s e n t e d  (see Van W i n k l e always  the  (Dalke  description.  I have used  and Sime 1938) both  techniques  deer  developed  the s t a t i s t i c a l  deer  locations.  can  be t r e a t e d l i k e  the  variability  Thus, of  in  univariate  2.9.  Track  movements. central  often  these techniques, any o t h e r  about t h a t  statistics  and  mean  although spatial  minimum home spatial  the  minimum  cores  of  home  Geographical  f o r analysis of  the l o c a t i o n s of a  variable,  deer  and a mean l o c a t i o n  location  can  be  calculated.  home r a n g e d a t a , I p r o v i d e  c o m p a r a b l e t o t h a t commonly  a  level  available  data.  winter  and  early  spring,  night  counts  were n o t  a l w a y s p o s s i b l e i n h i g h e l e v a t i o n s . I n o r d e r t o d e t e c t when were i n t h e s e  crossed be  spring  areas,  t r a c k counts  were made. Mount C a i n Road  and t h e h i g h e s t e l e v a t i o n  or followed along the road  used  to  and  Counts  During  travelled  The  The minimum home  located.  (1966) were a d a p t e d  t h e a n a l y s i s o f my  descriptive  for  Using  most  by N e f t  o f ways.  does p r o v i d e a simple  identify  were  considered  interpreted  hand, t h e t r a d i t i o n a l  r a n g e method was m o d i f i e d t o where  usually  i n a number  easily  r a n g e methods t o d e s c r i b e d e e r  range  are  (1975) f o r r e v i e w ) ,  provide  other  r a n g e method  home  o f animals  determine  and when d e e r  when d e e r  last  a t which a d e e r  track  leave these  visit  high  was  either  was r e c o r d e d . T h e s e d a t a first  deer  can  elevations i n  e l e v a t i o n s i n winter.  20  2. 10.  Night  Counts  P o r t i o n s o f the study spotlight road  shone  from  area  a vehicle travelling  t r a n s e c t s . Three n i g h t  weeks.  Because  sometimes intense  of  made  relocation  conducted fixed  counts  weather  less  of  population. reflective night.  provide  window-mounted  2.11.  During  certain  deer,  on  and f l a p  tags  tags  these deer  These  night  of t h e deer  were c o v e r e d  with  when i l l u m i n a t e d  usually  20 t o 45 power s p o t t i n g  of were  relative  movements  visible  could  of  counts along  the year.  measure o f s e a s o n a l  the  night  sighted  throughout  periods  be r e a d  at  through  a  telescope.  Sample S i z e Funding,  of  attempted  estimates  and s o were r e a d i l y  Numbers  predetermined  were  Badio-transmitters tape  a  and o t h e r commitments, c o u n t s  abundance.in the cutover areas a r e an i n d i r e c t  along  using  two  radio-tagged  transects  at night  every  frequently.  were  each n i g h t . The numbers o f d e e r  road  counts  were s e a r c h e d  study  deer.  One  failed yearling  logistics  animals  to hunters  limited  and  early  i n the study.  male e m i g r a t e d . i n the study  hunting  was  of  the  e i g h t deer  area a t t h e opening  three of the remaining  i n the study  eight radio-tagged  tagged  in  A n o t h e r t r a n s m i t t e r was l o s t  This l e f t  not r e s t r i c t e d  losses  t h e number o f r a d i o - t a g g e d  o f t h e 10 r a d i o - t r a n s m i t t e r s a t t a c h e d t o d e e r  radios  One  o f c a p t u r i n g and m o n i t o r i n g ,  deer  was  killed  with  of hunting area deer early  when a  functioning season.  during were  1975  1975  Deer and  shot.  i n September and  s i n c e I h a d o n l y a few l o c a t i o n s f o r him I d i d n o t i n c l u d e  them  in  were  the  seasonal  movements  analysis.  The two o t h e r d e e r  21  killed deer  i n early were  monitored consists and  of  for  a  in  of t h e complete  broad  This  t h e y were n o t study  thus deer  deer.  of the r a d i o - t a g g e d deer a r e r e p r e s e n t a t i v e occur  and  indicate  the p o p u l a t i o n . Track  counts  the radio-tagged deer, track  p r o v i d e an a c c u r a t e d e s c r i p t i o n  black-tailed  cycle.  These  s e a s o n a l movement p a t t e r n s o f f i v e  trends i n the seasonal d i s p e r s i o n  Combined, should  of  seasonal  p a t t e r n s o f two  movements  observed.  t h e a n a l y s i s e v e n though  complete  movements t h a t c a n  patterns  of  included  the p a r t i a l The  w i n t e r and some movements were  deer i n t h e study  area.  potential  movement  and n i g h t c o u n t s  o f the deer  give  population.  counts, and n i g h t  counts  o f t h e s e a s o n a l movements  22  3.  Results  3.1.  Snowpack I n The  Study  E l e v a t i o n s below October  1975  i n O c t o b e r and  900  m).  of  300  ground  during  the  before  melting.  snow  and  m by  first  at  This accumulation maximum  m)  few  in  increase  of  is  partly  and  the  of  289  between and  depth  south  and  west above  1000  Mid-elevations slightly  later  below  cm and  than  few  was  1200  m.  The below  (700  t o 900  did high  m)  mild  r a t e o f snowmelt  and  the  late 1200  melts,  accumulate.  March m.  In  when  April,  melted  by  a the  early  2 shows a s u b s t a n t i a l  of the  stations 1000  the days  This difference i n  depth  Mount C a i n  were  m and  depth  of  received  on  between  their  Road  aspects southwest  first  the  e l e v a t i o n s , except i s g r e a t e r than  that  the  snowpack d e c r e a s e s  periods.  The  at  t i m e between e a r l y December and  any  on  snow to  at  snow  e l e v a t i o n s . They have a p a t t e r n  s i m i l a r t o high  snowfall  m.  lower  m.  snow a c c u m u l a t i o n weather, the  Figure  to  bottoms  and  m began  at  (above  f o r o n l y a few  snowfalls  until  late  occurred  snow d e p o s i t e d  lasted  o f a change i n a s p e c t  southwest  valley  completely  1000  stations.  m  elevations  the  occurred  snowpack above  a result  snow  snowfalls  snow c o n t i n u e d  areas  i n the  reached  e l e v a t i o n s above 900  depth  most  high  1200  p r o g r e s s i v e l y extended  first  snowpack began t o d e c r e a s e July  to  mid-November. The  A f t e r the  deposited  s n o w f a l l s below  snowfalls  (below 700  elevations  first  were r e s t r i c t e d  Subsequent  elevations  m were s n o w - f r e e f r o m J u l y u n t i l  ( F i g . 2 ) . The  late  snow  1200  Area  snowpack i n m i d - e l e v a t i o n s  d i d not  during  rate for  of  brief  completely  e a r l y May.  of  Snow d i d  melt not  23  Figure along  2.  Perspective plot  Mount  Cain road.  of d a t e ,  elevation  Most w i n t e r r a n g e s  snow d e p t h s were s h a l l o w  and snow c o v e r  ranges  had  above  continuous  700  m  snow c o v e r o v e r  deep  winter.  and snowpack  were below  700 m  was i n t e r m i t t e n t .  accumulating  depth where Summer  snowpacks  and  25  accumulate period  i n low e l e v a t i o n s  or  to  elevations. free. than  as  great  were  depth,  snow-free  were h i g h e l e v a t i o n s Mount  increasing  Cain,  elevation  persistent  snow  after  of  except  of the difference  the  sidehill  is  a  Snow d e p t h s  floor  result in  of  Croman  they  than near  1976, much  sooner  snowpack  increased  had  floor  with  along the  deeper  and  more  were  their Valley  on  Mount  differences were  not  Cain.  floor  and  i n s l o p e and monitored  as  However, snow  was  l o n g e r i n upstream  parts of  t h e mouth.  Abundance Of Deer Food Important  area  are  species,  food p l a n t s o f b l a c k - t a i l e d  Alectoria  sarmentosa  Rubus s p e c i e s , C o r n u s  Epilobium  Pseudotsuga  menziesii  (Mirb.)  Besides tailed  these deer  the  palatable  species  L.,  Sarg.  of  (Jones other  black-tailed  shallon  L.,  Thuja  deer  the  study  ,  Vaceinium  Linnaea  borealis  (L.)  plicata  Roth.,  Donn,  p l a n t s a r e e a t e n by b l a c k -  and  1968). the  The  diversity  predominance o f  s p e c i e s a l l o w e d e s t i m a t e s t o be made o f t h e amount  available  a t each  habitat  and  1975, R o c h e l l e i n ££§_£._).  (Cowan 1945, Brown'1961, G a t e s  diet  in  Blechnum s p i c a n t  Franco,  many  deer  , Gaultheria canadensis  angustifolium  Tsuqa h e t e r o p h y l l a (Raf.)  food  April  snow-  i n snowpack between t h e v a l l e y  f r e g u e n t , and p e r s i s t e d  in  late  frequently  deeper,  !•#  a  were  f o r on t h e v a l l e y  as  more  long  a s i t d i d i n h i g h and m i d -  the  intensively  3.2-  as  cover than t h e adjacent s o u t h - f a c i n g s i d e h i l l .  Part  the v a l l e y  for  (Fig. 2).  depth  D a v i e R i v e r . The v a l l e y  aspect.  a  m)  S e v e r a l t i m e s i n w i n t e r , low e l e v a t i o n s  They  On  (300 m t o 700  assessment  site  and a  of  composite  26  summary o f f o o d r e s o u r c e s a v a i l a b l e and  serai  stage  was  therefore  these h a b i t a t s consisted shrubs,  herbs,  component  was  and  of  using  each  plant  calculated.  four  conifers.  estimated  in  major  The  Food  resources  components;  relative  differing  association  lichens,  contribution  techniques  as  of  of  each  explained  below.  Lichen The in  the  deer  percent cover of l i c h e n ,  forest is  lichen obtained  estimates  the  =  2  about  it  was  (1978).  to  log,  different  the  12%  percent and  0  zero  the  (p <  thus  1978). F o r  likely  to  amount o f f o o d  total  s i n c e the  cover  contribution  small, t h i s  total  amount  annual  lichen  overestimate of  has  the  available  litter-fall  between  in  food. high  biomass  the  two  of determination i s regression  purpose  line  is  that f a l l s  as  deer  of t h i s  food,  analysis, summer.  the c o n t r i b u t i o n of  lichen  a  occurs over  to  the  to the only  and  .06).  available  of l i c h e n  of  a linear regression  i s available  overestimates  cover  independently  standing crop of l i c h e n  w i n t e r and  (Stevenson  of  assumed t h a t s i m i l a r l i t t e r - f a l l  the  were  coefficient  slope  from  p r o p o r t i o n of the the  the  T h i s assumption  is  the  ,  radio-tagged  where p e r c e n t  estimates  Both  transformed  0.41),  ground over  is  some s i t e s  biomass  ( F i g . 3). although  statistically The  I . On  sarmentosa  by  t o o b t a i n an e m p i r i c a l r e l a t i o n s h i p  variables (r  Stevenson were  performed  low  measured,  by  Meetoria  c a n o p y o f s e a s o n a l h a b i t a t s used  shown i n T a b l e  was  mainly  d u r i n g summer. However, available slight  Krumlik's elevation  f o o d i n summer effect  on  the  (1979) m e a s u r e s o f forest-types  on  27  Table  I. Percent  forested  plant  radio-tagged total  of  during food  Forested  Cabbage,  SC = Sword  Western  Hemlock,  AW = A m a b i l i s  and e s t i m a t e d  associations deer  weight  indicated.  cover  a v a i l a b l e i n summer  and immature s e r a i the year.  available  plant  food  stages  used  Mean and s t a n d a r d  in  associations  each :  habitat  by  error of type  VS = V a c c i n i u m  are  - Skunk  F e r n - W e s t e r n Red C e d a r , DW = Deer F e r n WP = W e s t e r n  F i r - Western  Hemlock -  Hemlock,  SD = S a l a l  Hemlock - C o p p e r b u s h .  N = Newly L o g g e d , H = Herb, F = F e r n ,  Immature  -  Plagiothecium, -  Douglas-fir,  SW = S a l a l - W e s t e r n Hemlock, AT = A m a b i l i s F i r - T w i s t e d MC = M o u n t a i n  on  Serai  S = Shrub, C =  Stalk,  Stages Conifer.  :  Association or S e r a i Stage  Shrub  Herb Plot No.  % cover  kg ha~^  Conifer  kg ha""  cover  kg ha  Lichen cover  Total  kg ha  kg ha  VS  7  9.2  40.9  1005.7  26.0  79.6  17.8  11.0  1137.2  vs  20  20.9  93.0  700.7  26.0  79.6  28.4  76.4  949.7  VS  41  58.4  259.9  156.8  9.5  29.1  15.1  5.6  451.4  n=3 sc  13  2.1  9.3  10.5  8.0  24.5  25.6  19  2.6  11.6  18.0  22.0  67.3  17.6  10.5  n=1  94.0 107.4 107.4  WP  3  .4  1.8  .1  5.0  15.3  21.4  23.6  40.8  WP  39  1.3  5.8  1.4  5.0  15.3  21.3  23.2  45.7  n=2  43.2  i  23.2  122.0  9.3  110.8  AW  10  1.5  6.7  6.4  28.0  85.7  21.3  AW  18  1.0  4.4  5.3  30.0  91.8  17.1  AW  44  .9  4.0  269.7  7.0  21.4  32.6  135.6  430.7  AW  45  1.6  7.1  159.8  12.5  38.2  22.0  26.5  231.6  n=4  354.4  94.0  49.7  n=1  DW  t  846.1  223.8  1  3.5  148.3  Table I  (cont.)  Association or S e r a i Stage  Herb Plot No.  % cover  Shrub  _ kg ha  1  -  1  kg ha  L: hen  Conifer  -1  cover  kg ha  15.3  22.6  29.6  407.0  .0  23.2  33.0  680.9  SD  2  .6  2.7  359.  <f  5.0  SD  5  .9  4.0  643.9  .0  cover  Total  kg ha  n=2  kg ha  504.4  t  SW  4  .7  3.1  218.0  23.0  70.4  35.9  201.9  ^93.^  sw  6  1.1  ^.9  218.1  10.0  30.6  23.9  37.^  291.0  sw.  8  1.4  6.2  253.6  8.0  24.5  20.9  21.4  305.7  sw  17  .7  3.1  21.4  28.5  87.2  17.7  10.7  122.4  sw  46  15.3  28.5  77.5  261.6  164.4  1.0  5.0  n=5  294.8  i  137 .7  132 .7  AT  1  2.4  10.7  323.5  12.0  36.7  31.8  122.1  493.0  AT  28  1.4  6.2  1018.0  26.0  79.6  32.0  125.3  1229.1  AT  29  1.*f  6.2  16.0  49.0  24.9  44.3  378.9  279. ^  n=3  MC  32  .7  3.1  406.5  6.0  18.4  MC  33  12.5  55.6  531.0  1.0  3.1  13.1  18.2  700.3  -  ^ 6 1 .5  3.1  431.1  12.1  601.8  n=2  516.4  t  120.7  Table I  (cont.)  Plot No.  N  38  of /O  cover  1.2  Conifer  Shrub  Herb  Association or S e r a i Stage  kg ha  -1  5.3  kg ha""  1  5.4  cover  33.5  Lichen  kg ha  102.5  cover  Total  kg ha  .0  .0  9  113.2 113.2  n=1  H  kg ha  41.0  182.9  5.7  .5  1.5  .0  .0  190.1  H  11  36.8  163.8  .3  .0  .0  .0  .0  164.1  H  14  61.3  272.8  .3  2.5  7.6  .0  .0  280.7  H  15  42.1  187.3  .5  1.0  3.1  .0  .0  190.9  H  16  13.1  .0  3.0  9.2  .0  .0  67.5  H  21  56.5  251.4  20.1  10.0  30.6  .0  .0  302.1  H  23  36.2  161.1  83.8  6.5  19.9  .0  .0  264.8  II  24  91.2  405.8  12.1  4.0  12.2  .0  .0  430.1  H  30  11.5  51.2  55.5  1.0  3.1  .0  .0  109.8  H  31  32.2  143.3  3.4  1.5  4.6  .0  .0  151.3  58.3  n=10  F  12  43.9  194.5  .0  8.5  26.0  .0  .0 n=1  215.1  -  220.5 220.5  106.1  Table I  (cont.) Total  S  27  1.9  8.4  234.0  21.0  64.3  .0  .0  306.7  S  34  12.3  54.7  39.6  4.0  12.2  .0  .0  106.5  S  36  31.6  140.6  .2  2.5  7.6  .0  .0  148.4  S  37  38.9  173.1  7.5  8.5  26.0  .0  .0  206.6  S  40  84.6  376.5  3.0  23.0  70.4  .0  .0  449.9  S  43  42.5  189.1  290.0  6.5  19.9  .0  .0  499.0  cover  Shrub  Lichen  Plot No.  of  Herb  Conifer  Association or S e r a i Stage  kg ha  _1  kg ha  -1  cover  kg ha  -1  of /o  cover  kg ha  -1  kg h a "  286.2  n=6  t  12.7  6o.o  183.6  .0  .0  315.1  469.4  28.0  85.7  .0  .0  584.9  8.4  461.2  80.0  244.8  .0  .0  714.4  - 3  5.8  515.8  59.5  182.1  .0  .0  703.7  6.0  26.7  141.3  115.5  353.4  .0  .0  521.4  118.8  C  22  26.7  C  25  6.7  29.8  C  26  1.9  C  35  1  c  42  n=5  567.9  -  1  161.3  163.0  32  Figure  3. L i n e a r  biomass  regression  transformed  L = .0006 P * - i s  t  lichen  Lichen  cover.  increases  by  where  i n percent  of percent  L  log is  biomass  cover  of  l o  cover  .  lichen  r  2  on  = .11,  lichen p<  .06.  b i o m a s s and P i s p e r c e n t  increases  lichen.  of l i c h e n  exponentially  with  2 0  P  E  R  C  3 0  E  N  T  C O V E R  O F  L I C H E N  34  coastal this  British  study high  Fir  -  annual  Stalk  study  and  were e s t i m a t e d  lichen  my  plant  to  and  426  Hemlock  estimates  ha  similarity  l e d me  .  In  Copperbush  376  kg  between  to accept  - 1  (Amabilis  -  have between 9 and The  kg  associations  Mountain  litter-fall.  (1979) measures and my  r a n g e d between 71  elevation forest  Twisted  associations) of  Columbia  the  ha  - 1  Krumlik's values  for  area.  Shrubs Almost Vaceinium The  a l l shrubs at parvifolium  c o n t r i b u t i o n by  estimated three  as  increasing  the  stem  variation  i n the  variation  in  coefficient  of  variation heights.  of the  to the  vertical  height  from  rate  current  year's (Fig.  stem h e i g h t annual  annual  available  a n n u a l growth 4) .  In  is  to  deer  growth b e i n g  coefficient  more p r o s t r a t e  of  with  explained  only by  g r o u n d t o crown t o p  may  not  be  the  shallon  significantly  of  the  in  stem  likely  the  , the  rate  different  with of  from  compared Hence,  predictor  . In both Vaceinium  i n annual growth i n c r e a s e s  In G a u l t h e r i a  the  species. best  species,  ,  shallon  Vaceinium  with  the  49%  is  was  of  variation  determination  the  more shallon  g r o w t h f o r m o f G.  growth forms o f  growth i s n o t  or  .  In a l l  increased  Gaultheria  lower  were  shallon  In both Vaceinium  a c c o u n t s f o r 72%  growth.  determination  of i n c r e a s e  stem h e i g h t .  sites  , or G a u l t h e r i a  food  a n n u a l growth i n G a u l t h e r i a s h a l l o n the  Assessment  a n n u a l shoot growth of t h e s e s p e c i e s .  i n annual  result  , Xs. a l a s k a e n s e  height  T h i s low  Habitat  shrubs to the  the  species,  the  species,  increasing  increase one  of  (p>.05).  in  35  Figure.4. in  Height  shrubs  of shoot annual  important  H = h e i g h t from  as  t h e ground  Vaccinium  p a r v i f olium  :  Vaccinium  alaskaense  :  Gaultheria  shallon  :  deer  growth  p e r stem and stem h e i g h t  f o r a g e . ft = a n n u a l  t o the top o f t h e shrub  growth ( g ) ,  crown  (cm) .  A = .00056 H*- *••<>, p<.00, r = . 85 A = .0093 H - 3 2 1 . 3 0 , p<.00, r = . 7 2 A = .019 H » - p < - 0 0 , r=.49 8 8  L  2  2  2  37  The three  number  shrub  sites.  species  Using they  produced  by  shrubs  the  of  Habitat  cm  the  at  less  growth of  stems  annual  sites.  s h r u b s was  cm  high.  assigned  sites,  contribution to t o t a l  cm  s m a l l and  was  difference there  to  heights  use the  depending  less  30  than  cm.  the  the  of  short  assigned  height  t h o u g h t t o be  calculations  upon  to  were the  counted  as  very total of  but  not  a height  height  was  of at  chosen  30  total  stems l e s s  cm  (Appendix  growth by  30  1 ) . At  cm most  than  makes  30  little  a n n u a l g r o w t h . . I n some total  of  annual  than  stems l e s s  height  sites,  annual  growth  assigned  height chosen f o r  stems  were m a i n l y  i n h a b i t a t s where  there  stems of  10  of cm  c o n t r i b u t e d by  Gaultheria  shallon  f o r stems l e s s than  most r e p r e s e n t a t i v e and  of food  were  average.annual growth  assigned of  high  comparison, the  annual  estimate  These s i t e s  high d e n s i t i e s  Overall was  the  and  either  cm  were a s s i g n e d  were s u b s t a n t i a l d i f f e r e n c e s i n t h e  estimate  are  of  these  g r o w t h of s h r u b s p r o d u c e d  For  of 0 cm  In  increase e f f i c i e n c y  also c a l c u l a t e d with  being  growth  parvifolium  little  This assigned  i t best represented 30  30  were  high  annual  and  growth r e l a t i o n s h i p  To  stems cm  V.  than  shrubs.  30  of the  the  assessment  estimated.  contributed  l e s s than  than  Habitat  than  less  these  Assessment  because I f e l t  was  same h e i g h t - a n n u a l  Assessment  Habitat  site  a v a i l a b l e from  Stems  the  for  g r o w t h r e l a t i o n s h i p s ( F i g . 4)  species other  sites  in calculation  stems  within  each  Shrub  a t most  food  measured. 10  .  stems were d e t e r m i n e d  same f o r a l l h a b i t a t t y p e s ,  t o have t h e  numerous b u t  of  plots  Vaceinium  alaskaense  amount  on  were t h e  calculations,  V.  heights  the height-annual  assuming  considered  and  shrubs  so  was  used  (Table I ) .  in  30  cm a l l  38  Herbs The  contribution  available  at the  converting ha-*).  cover  The t o t a l  cover  of  percent ,  were  is  was c o n v e r t e d  (1968).  I n Gates  percent  cover  covers  )  a percent  t o biomass  herbs  except  i n T a b l e I . The t o t a l  percent  o f a l l h e r b and f e r n  species  and  Species  cover  from  the f l o r i s t i c  with  l e s s than  o f 0.1. The t o t a l calculated were  of annual  - 1  of  cover  covers  was o b t a i n e d .  forage  for  conversion  factor  comprising  the  1%  depends  percent  from  Gates  at  which  given  herb p r o d u c t i o n  cover  upon  cover  biomass  the  as  of forage  gave  4.45  heights  estimate  of  (Traczyk  the  made  food  Epilobium  anqustifolium  percent  cover.  mitigate  t h e problem  between  species.  of  This  in  was  the  predominance of d i f f e r i n g  to  the  species of  biomass  total with  the  greatest  E. a n q u s t i f o l i u m : cover  Table  estimation  I, of  acknowledging  herbaceous  would  relationships  potential  forage  herbs source,  The c o n t r i b u t i o n o f h e r b s t o t h e t o t a l  food i s given i n  involved  contribution  herbs  and T r a c z y k where  substantial  kg  o f t h e h e r b l a y e r . The  1977). However, i n most o f t h e h a b i t a t a s s e s s m e n t s a  35%  d i v i d e d by  and a mean d r y w e i g h t This calculation  5%  percent  b i o m a s s was e s t i m a t e d  ground  by  ferns,  (1968 : 4 9 ) , f o u r s i t e s  r e s p e c t i v e percent  per p e r c e n t ha  estimated  i t s wet w e i g h t . The d r y w e i g h t s o f h e r b s were t h e n  their  food  f e r n s t o b i o m a s s (kg  by a f a c t o r  and wet w e i g h t  t h e deer  was  and was o b t a i n e d  were m e a s u r e d . Dry w e i g h t o f t h i s of  and  of  given  to  sites  Assessment s i t e .  assigned  ferns  herbs  cover  P*. a q u i l i n u m  a t each H a b i t a t  and  Assessment  i s t h e sum o f p e r c e n t  cover  herbs  cover  aquilinum  (excluding list  Habitat  percent  Pteridium  of  using  amount errors  t h e above  39  method.  Conifers The the  c o n t r i b u t i o n by  Habitat  c o n i f e r s to the  Assessment s i t e s  was  to t h a t used f o r h e r b s .  Percent  differently  and  the  in  availability  areas,  the  conifers  to convert calculated  49)  o f the by  three  to.the  percent  cover  wet  Gates  measured  sites.  decimal  point  conversion conversion equalled  and  so  factor.  3.06  as  was Using  - 1  were  10 m h i g h  had  areas, c o n i f e r s  branches.  A  factor deer  the  percent  cover  deer  forage.  At one  not  of c o n i f e r  the trend  t o be used  the  was,  forage.  of  in  from of a  of  : and  sites,  the  other  misplaced  calculation  1%  (1968  conifers  three  as  menziesii  of these  a result  remaining I  was  w e i g h t s were e x p r e s s e d Pseudotsuqa  2  used  food  were g i v e n i n G a t e s  appeared  of  approaches  sites  than  was  cover  for  used i n Table  kg h a  two  logged  10 m h i g h  t o biomass o f  dry  similar  the  data  much h i g h e r  This value  factor  The  Four both  available  b i o m a s s was  reaching  at  estimated  percent  ground. In unlogged  using  In  c o n i f e r s below  of c o n i f e r s  (1968).  the  u s e d . The  food  to d i f f e r e n c e s i n  deer.  areas, was  (1968).  weights  for  due  was  of c o n i f e r s below  areas,  have low  from Gates  biomass  conifer  2 m high  d i d not  a t which he  their  cover  forested  than  reaching  10 m h i g h  given  In  food  deer  i n a manner  of c o n i f e r s  unlogged s i t e s  because, i n logged  branches  42%  I.  less  employed  percent  estimated  cover  of c o n i f e r s a s  total  used i n T a b l e  to  logged  available  of  the  sites,  the  conifer  cover  40  3.2.1.  Abundance Of Food D u r i n g  Total  amounts  given i n Table available intervals  of  of food a v a i l a b l e  I. Estimates  t o deer  in  t o deer  made by G a t e s  summer  my e s t i m a t e s .  Coastal Douglas-Fir  Summer  are  Gates  a s s u m p t i o n s t h a t I made t o e s t i m a t e  Average  food  95%  i n Figure  5. F o u r o f t h e n i n e  (1968) worked i n t h e  shrub  the  other  five  most o f t h e a v a i l a b l e  Amabilis  Western again  Hemlock,  f o o d was c o n t r i b u t e d by  t h e main  Sword  had  moderate  component.  Fern-Western  Red  remaining  Cedar,  these  plant  association, the  Hemlock  three  in  Conifers  one  mainly  Salal-  shrubs  were  associations,  the  food.  of c o n i f e r s .  Vaccinium-Skunk  portion of t o t a l  total  (a  food  abundance  Food In a l l  Cabbage  available  (15%),  i n any o f t h e o t h e r  as abundant a s h e r b s i n young  provided  s e r a i stage.  site  to  abundance t h a n  about t w o - t h i r d s  logged  and  food.  V a c c i n i u m - S k u n k Cabbage a s s o c i a t i o n , h e r b s were a m i n o r  component compared greater  except  h e r b s were a s m a l l  abundance  D e e r F e r n - W e s t e r n Hemlock, and  a s s o c i a t i o n s was c o m p r i s e d  associations,  Stalk,  shrubs.  Western H e m l o c k - P l a g i o t h e c i u m , had s m a l l amounts o f  In  (Vaccinium-  four associations  amounts o f f o r a g e ;  The  stage i s  Fir-Twisted  a s s o c i a t i o n s . In these  data  productivity.  associations  Two a s s o c i a t i o n s , A m a b i l i s F i r - W e s t e r n  in  our  M o u n t a i n H e m l o c k - C c p p e r b u s h ) h a d much g r e a t e r f o o d  than  biomass  confidence  abundance f o r e a c h a s s o c i a t i o n and s e r a i  Skunk Cabbage, S a l a l - D o u g l a s - f i r , and  the  B i o g e o c l i m a t i c Zone, a g r e e m e n t o f  supports  presented  summer a r e  (1968) o f f o o d  within  Although  during  most o f t h e a v a i l a b l e  Data f o r t h i s  Salal-Western  serai  Hemlock  stage  but  were  a s s o c i a t i o n s and serai  food  in  stages. the  newly  were b a s e d on o n l y  a s s o c i a t i o n before  i t was  41  Figure  5. Abundance o f f o o d  habitats  used  by  assessment s i t e s plant  WP = W e s t e r n  radio-tagged  are indicated  associations  Fern - Western  a v a i l a b l e t o deer  :  Red  above  VS = V a c e i n i u m Cedar,  over the  Hemlock - C o p p e r b u s h  ..  Number o f  histograms.  Forested  F i r - Twisted Serai  Logged, H = Herb, F = F e r n , S = Shrub, C =  - Western  AW = A m a b i l i s  - Douglas-fir,  Immature  in  the year.  Fern  Hemlock - P l a g i o t h e c i u m ,  AT = A m a b i l i s  summer  - Skunk Cabbage, SC = Sword  DW = Deer  W e s t e r n Hemlock, SD = S a l a l Hemlock,  deer  during  SW = S a l a l Stalk, Stages  Hemlock, Fir - Western  MC = M o u n t a i n :  Conifer.  N = Newly  herb |  80CH  | shrub  lllllll  conifer  iHH  lichen  60CH  Q O  o  400-j  LL_  200H  -r S  -T"  VS  SC  DW  Coastal  WP  Western  AW  Hemlock  SD  SW  AT  MC  Mountain  Zone  Hemlock  Zone Forested  Plant  Associations  N Immature  Serai  Stages  43  logged).  At  other  associations, available  food.  s t a g e had l e s s  In  herb,  did  herbs  food  was  were  more  abundance i n t h e c o n i f e r serai  replaced  stages  contrast  consisted  forested  total  available  (Fig.  herbs  food  fern  serai serai  abundance due t o serai  stage  by s h r u b s and c o n i f e r s a s t h e  than  to forested  plant  In  this  serai  stage  conifer food. Total  plant  mainly o f herbs. associations  less 5) .  abundance  food  abundance,  i n summer than The to  l e s s abundant  than  associations  was s h r u b s , f o o d  than  in  food other  some  young  serai  some f o r e s t e d  forested  stage plant  Fir-Twisted  plant  serai  most  plant  was  Stalk  in  greater  had more  associations plant  the  .  food ,but  associations  comparable  associations  of  associations  stages  forested  i n t h i s s e r a i stage than  Cabbage and A m a b i l i s  shrubs comprised  did forested  serai  i n young  s e r a i s t a g e s h a d much  t h e most p r o d u c t i v e conifer  where t h e main  O l d s e r a i s t a g e s were s i m i l a r  i n that  a v a i l a b l e f o o d . However, o l d e r  much  and  s e r a i s t a g e was g r e a t e r  amounts o f c o n i f e r f o r a g e t h a n In  component o f  In the o l d e s t  food.  abundant  herb  total  ( F i g . 5).  component o f t h e a v a i l a b l e f o o d  the  plant  stages. In  to  the greatest  the  i t had a g r e a t e r  m a j o r components of t h e a v a i l a b l e shrub  climax  f e r n , and s h r u b s e r a i s t a g e s ,  than  amounts o f s h r u b s  considered,  different  component o f a v a i l a b l e f o o d . The s h r u b  herb f o o d  however,  increased  with  h e r b s o r s h r u b s may f o r m  formed t h e g r e a t e s t  stages;  sites  in  , but food  food was  Vaccinium-Skunk  associations.  44  3.2.2.  abundance Of  During  Food D u r i n g  winter,  Winter  snow a c c u m u l a t i o n  and  t h e amount o f f o o d a v a i l a b l e  to  availability  in  i t can bury  restrict  deer  mobility  exposed  food.  The be  degree  two  ways: and  deer.  thus  Snow  e v a l u a t e d by c o n s i d e r i n g t h e  vertical  growth of s h r u b s  major  winter  total  stem  (Fig.  6)  height  on  plots  t h e amount o f f o o d the  height-annual  that  the  depth,  growth  w i t h i n each  be  simulated  above  a  could  be  shrubs  of  for  the  (37  then  at the  growth  the  1975,  by  crown  species  i n F i g u r e 7.  amounts  food  each  stem  amount  stem can  shoot  calculated  so t h a t  Such  of  snow  by  stems  Noting  evenly  or serai growth the  snow c o n d i t i o n s ,  over annual  stage  effect  of  snow  depression  would  of  i s not  cause  l e s s than  downward  can  occurring  s h a l l o w e r snow d e p t h s ,  above t h e snow t o be  from  assuming  of shrub  through  displacement  that  determined  distribution  a  eguations  shrub  i s apportioned  annual  is  proportion of  of  be  the  considered  ( F i g . 4 ) , and  of  burial  exposed  depths  plant association  effect  their  Under c e r t a i n  of  Rochelle in  (Fig. 6). Regression  vertical  accommodated  indicated.  shrub  relationships  forested  e v a l u a t e d . The  of  (Jones  H a b i t a t Assessment s i t e s .  produced  ( F i g . 7 ) . The  thus  three  predict  g i v e n h e i g h t was  and  deer  t o 49%)  a n n u a l g r o w t h o f each  crown  obtaining  i s c o n t a i n e d w i t h i n the crown and  source  to  food  current year's  s t u d y a r e a were a n e a r l y c o n s t a n t  were u s e d  measured  from  distribution  annual  i n the  affect  b u r i e s f o o d p l a n t s can  component o f t h e f o o d r e s o u r c e s . The  important  affect  s o u r c e s o r i t can  deer  shrub  p r e p . ) . Crown d e p t h s  ( F i g . 2)  can  food  prevent  t o which snow c o v e r s and  food  melt  the those  displacement  45  Figure  6.  Crown  d e e r f o r a g e . The height of  and  stem  relationship  height i n shrubs important  between  crown  i n shrubs a l l o w s p r e d i c t i o n s of the  shoot  ground  depth  annual  growth.  t o t o p c f crown  C = crown d e p t h  depth  vertical  and  as  stem  distribution  (cm), H = h e i g h t f r o m  (cm) .  Vaccinium  parvifolium  : C = -.70  + .49±.09  H , p<  - 0 0 , r =  .79  Vaccinium  alaskaense  : C =  1.51  • .37±.04  H , p<  . 0 0 , r =  .69  : C =  .97  + .49±.06  H , p<  . 0 0 , r =  .88  Gaultheria  shallon  2  2  2  47  7.  Figure  forested  Vertical plant  associations  biomass a v a i l a b l e abscissa. available  This  above  forested  serai  = Western  the  growth  annual  from  shoot  approximately  in  stages. Ordinate i s the the  amount  displacement  :  Red  by  VS = V a c e i n i u m  ground  on  of food t h a t  growth o f  50% o f t h e  be  shrubs.  three  h e i g h t above  the  would  of  main  the ground. shrub  80% o f t h e s h r u b  Cedar,  DW  SD = S a l a l  AT = A m a b i l i s  Hemlock - C o p p e r b u s h H = Herb,  .  = Deer  Immature  snowpack.  Fern AW  - Douglas-fir,  F i r - Twisted  F = Fern,  the  - Skunk Cabbage,  Hemlock - P l a g i o t h e c i u m ,  W e s t e r n Hemlock,  Logged,  height  annual  food  food i n  s t a g e s i s b u r i e d . T h e s e e s t i m a t e s do n o t i n c l u d e  associations  Hemlock,  serai  with i n c r e a s i n g  a r e 50 cm,  o f shrub  Fern - Western WP  shrub  h a b i t a t s i s b u r i e d , whereas  the e f f e c t s plant  available  s p e c i e s decrease  immature  is  and  of  above t h e snowpack a s s u m i n g no d i s p l a c e m e n t  When snowdepths in  the  biomass  E s t i m a t e s of t o t a l shrub  distribution  Serai  Forested SC =  - Western  Hemlock,  = Amabilis  Fir -  SW = S a l a l Stalk, Stages  MC :  S = Shrub, C = C o n i f e r .  Sword  Western  = Mountain N = Newly  k8  49  of  shrub  shallow  the complete  magnitude o f d i f f e r e n c e s  associations  comparison in  c o u l d permit  these  and  serai  of the s e n s i t i v i t y  t o snow d e p t h  distribution the t o t a l  in  shrub  o f shrub  abundance t o  of  shrub  annual  evident  available  f o o d abundance was l e s s  taller  same  needed  to  depth  of shrub  the  vertical  during  snow-free  periods.  a f f e c t e d by snow i n f o r e s t e d  than i n c u t o v e r s . I f only b u r i a l  by snow  75%  a r e a s snow d e p t h s  of  results  stems  snow d e p t h s of  is  accounted  for,  o f between 50 and 110 cm  the food  o f between  Of Snow On P o s t u r e  In nearly  provided  20 and  50  by s h r u b s ; i n  cm  achieve  the  the  Amabilis F i r - Twisted  a l l the shrub  1976  Of S h r u b s  and  Stalk association  stems were p r o s t r a t e on t h e ground  covered  by up t o 30 cm o f snow. At t h i s  mean d i s t a n c e and 95% c o n f i d e n c e l i m i t markers  on  Vaccinium  was 8.8 ± 4.2 cm distance melted the  snow  shrubs  habitats, cover  when  direct  effect.  Effect  6,  i n forests  i n forested  logged  between  because  c o n s i d e r e d and no d i s p l a c e m e n t  are  by  growth i s c o n s i d e r e d r e l a t i v e t o  than i n cutover areas. T h i s d i f f e r e n c e  then,  abundance  s t a g e s does n o t permit a  i s more  amount o f s h r u b s  Generally,  is  of shrubs  habitats (Fig. 7). Differences i n sensitivity  availability  were  burial  snowpacks.  The plant  crowns  alaskaense  from  the  ground  stems b u r i e d b e n e a t h  (n = 1 7 ) ; f o r M e n z i e s i a f e r r u q i n e a  was 6.2 ± 3.1 cm  and t h e s h r u b s  mean d i s t a n c e f r o m  (n = 3 ) . On J u l y  had r e t u r n e d t o t h e i r  2 after  a t 900 m, on  June  time the to  the  t h e snow  stems  this  t h e snow had  snow-free  postures,  t h e g r o u n d t o t h e m a r k e r s was 64.1 ± 14.5  50  cm  for  Vaceinium  ferruqinea.  Before  were d e p r e s s e d The  t o l e s s than  Vaceinium  were  t h a t deer  vertically  of  alaskaense  by  c r o w n s o f t h e stems  shrub On  crowns  June  6,  that  snowpack  V. a l a s k a e n s e  stems  and become e n t r a p p e d  are  stems  are  pulled  Where s h r u b s  when  their  encasing  the  as  The  by s h r u b s  but a l s o  snowpack Although  food  vertical  collapsing bottom sloping  process.  o f c r o w n s would  until  make  virtually  the  shrubs  a l l t h e snow had of  available  food  i s increased.  of shrubs  by snow i s d e p e n d e n t  on d u r a t i o n of snow  accumulation  and  melt,  cover, and  not q u a n t i f i e d , i n t e r c e p t i o n  Vaceinium  depth.  a r e c o v e r e d by snow i n w i n t e r and e n t r a p p e d i n  deer  burial  in  snow m e l t s . On  When t h e snowpack m e l t s , t h e amount  contributed  species  was  usually  the  n o t o n l y on snow  relative shrub's  of f a l l i n g insufficient  rates  of  structure.  snow by t o cause  crowns large  d i s p l a c e m e n t s o f t h e s t e m s . However, G a u l t h e r i a s h a l l o n  i n t e r c e p t s snow r e a d i l y after  with  by  Because  stems a r e e v e n t u a l l y r e l e a s e d a t t h e  t h e snowpack, t h e d i s p l a c e m e n t  depth,  down  buried  within i t .  s n o w c r e e p would a l s o c o n t r i b u t e t o t h e b u r i a l  melted.  of the  than  snowpack  unavailable  also  snow e v e n t h o u g h t h e snow was g e n e r a l l y l e s s  entrapped  terrain  was  most  snow m e t a m o r p h o s i s t h e snow c o m p a c t s and d e c r e a s e s  the  shrubs  and  snowpack. The s h r u b  of  the  snow-free h e i g h t . the  by  snow.  melted,  crowns were w i t h i n 10 cm o f t h e g r o u n d  deepening  The  of  near  food provided  30 cm d e e p . I t a p p e a r s the  14% of t h e i r  located  redistributed  with  and 102.0 ± 36.2 cm f o r M e n z i e s i a  t h e snow h a d c o m p l e t e l y  markers  suggesting  covered  alaskaense  a snowfall.  and c o n s e q u e n t l y  i s noticeably displaced  51  Amount Of The each  Food  A v a i l a b l e During  amount o f f o o d  s n o w f a l l and  o f snow d e p t h on 31,  1976  for  Winter  exposed above the  p e r i o d o f snow m e l t .  food  the  effect  a b u n d a n c e , snow d e p t h s r e c o r d e d  on  March  H a b i t a t Assessment s i t e .  used f o r l o g g e d estimated  areas.  using  with  To i l l u s t r a t e  were a p p l i e d t o t h e v e r t i c a l  each  snowpack changed  of shrub  Snow d e p t h s i n F i g u r e  Snow d e p t h s  the  distribution  in  relationship  forested modified  food  2 were  habitats  from  were  Fitzharris  (1975) :  C = -26.0  + 0.65  L  C = the  amount o f snow b e n e a t h  logged  areas.  water  Fitzharris  eguivalent  densities  inside  multiplied  the  amount  of  snowpack was and  did  were  not  winter  covered conifer  as  (Fig. to  forested  outside by  on  they  of  by  snow.  young Since  available  depth.  assumption  trends  i n food  h a b i t a t s , the  snow  and  the  would  areas not  conifer are  during  snow  stems.  Herbs snow  availability  assumption  would  component i n young  short only  by  these  same  the  and  would  be  a s m a l l amount snow-free  of  periods  make s u b s t a n t i a l d i f f e r e n c e s  abundance. I n conifer  during  This  was  burial  I  The  e x p o s e d above  of the  t o have t h e  trees  i n these  snow  estimate  deer  there  in  of  Therefore  summer. of  in  forested areas.  shrubs  to  depth  trends  b a s i s of d i r e c t  in  abundance  snow  snow d e p t h s i n cm  consistent  10 t o  the  had  where t h e  5), t h i s  annual  and  unavailable  the  with food  no  C o n i f e r s were t h o u g h t  stages  L =  i n c l u d e downward d i s p l a c e m e n t  overestimate serai  found  contributed  considered  conditions. in  but  calculated  canopy,  (1975) e x p r e s s e d  y intercept  food  a  older serai  understory  was  stages  and  much t a l l e r .  in  Snow  52  would c o v e r  some of t h e l o w e r  same  raise  time  conifer  food  sources  Consequently, estimation depths  the  five  total  lichen  within  March 31 of  component  and  of  others.  The  vertical  affected  i n areas while  downward  w i t h each  snowpack. On depths  be  s u b j e c t e d t o the On  the  elevation  occurred  which  than  e x c e p t i o n a l l the Stalk  were above 700 and  lower  shrub  on  m.  or The  of  the the  They  partial  depths  consider on  and  these  minor  in  would  be  most  reductions  in  the  snow c o v e r  throughout  progressively  displaced  metamorphosis o f  Mount  Cain  where  the snow  700-800 m  had  w i n t e r . These e l e v a t i o n s would of  more n o r t h e r l y  above  one  Fir-Twisted  m  shrubs  subsequent  throughout  snow  on  t h e snow  displacement  of  would be  displacement  would e x t e n d  With  678  of shrub  the s o u t h - f a c i n g s l o p e of  snow c o v e r  snowpack. the  12%  i n F i g u r e 2 were o b t a i n e d , e l e v a t i o n s above  continuous  of  as  above  a continuous  s n o w f a l l and  range based  f o r some h a b i t a t s  winter. In these a r e a s , shrubs  deer.  was  i n F i g u r e 8.  distribution  having  of  1978).  exposed  where t h e r e were still  the  estimated  are i l l u s t r a t e d  substantial  reach  lichens  (Stevenson  effects  the  previously unavailable  over  by  was  food  at  not i n t r o d u c e major e r r o r s i n  contribution  shrubs;  would be  be would  biomass present  burial  snowpack  The  estimates  on  estimates  would  month l i t t e r - f a l l  These recorded  that  conifer  encountered.  f o o d s o u r c e s , and  s u b s t r a t e so  the assumption  of  total  only  the  conifer  shrubs  facing  by  the  settling  and  less  steep s l o p e s ,  entrapment  and  displacement  700-800 sites  m.  classed  as e i t h e r  Mountain Hemlock-Copperbush exceptional site  was  at  an  amabilis  associations elevation  a n o r t h - n o r t h e a s t - f a c i n g s l o p e . A l l of the  of  sites  53  Figure  8.  Abundance  of food  available  to deer during  s e a s o n a l home r a n g e s . Number o f a s s e s s m e n t s i t e s above would  the  h i s t o g r a m s . Arrows i n d i c a t e t h a t  be r e d u c e d b e c a u s e o f  snow. Cabbage, Western  Forested  plant  SC = Sword Hemlock, Fir.-  displacement  associations  of  WP  = Western  = Amabilis  Western  SW  = S a l a l - W e s t e r n Hemlock,  MC  = Mountain  .  H = Herb, F = F e r n ,  component crowns  by  VS = V a c c i n i u m - Skunk DW  SD = S a l a l  AT = A m a M l i s  Hemlock - C o p p e r b u s h  their  Hemlock -  Hemlock,  on  indicated  the shrub  F e r n - Western Bed C e d a r ,  AW  N = Newly L o g g e d ,  :  are  winter  = Deer  Fern  -  Plagiothecium, -  Douglas-fir,  F i r - Twisted  Immature S e r a i S = Shrub, C =  Stalk,  Stages :  Conifer.  I |  I shrub 1 conifer llHU lichen  H SC  DW  Coastal  WP  Western  AW  Hemlock  SD  SW  AT  MC  Mountain  Zone  Hemlock  Zone Forested  Plant  Associations  N  r°h  T1  H  F  Immature  Serai  T  S Stages  55  are  w i t h i n the a l t i t u d i n a l  displacement likely  would  Another is  occur.  overestimated  Hemlock-Copperbush  Thus,  shrub  food  plant  association  abundance  Vaccinium-Skunk  Cabbage  Estimates  food  were b a s e d  t h r e e a r e a s i n t h e Croman V a l l e y Assessment  upper p o r t i o n s of the longer Davie  lasting  Valley,  e l e v a t i o n s on the  snow  Biver Valley.  Croman  deeper  of  the  f o r which f o o d  association.  Habitat  sites  valley.  abundance i n t h i s  were l o c a t e d  These s i t e s  had  than  did similar  Although  few  snow  snow  regime  was  the  cover  snowpack  and  shrub  Two  of  i n the middle  and  deeper  and  depths  were  similar  to that  displacement,  the  Vaccinium-Skunk Cabbage a s s o c i a t i o n  than  in Figure  winter  on  Amabilis  C o p p e r b u s h , and  Fir  Vaccinium  comparison  of  misleading.  Estimates f o r  based  on  comparable above was  the  and  higher  over-estimates of -  Twisted  Stalk,  - Skunk Cabbage  the below  with  food  in  amount o f would  mid-  food  be  less  abundance  associations,  winter,  1  in  plant  700  m and  displacement  associations  be  were  therefore should  of  -  direct  o t h e r s i n F i g u r e 8 may  other  t h e snowpack would be  elevations.  of  M o u n t a i n Hemlock  snowpack. S i n c e snow c o v e r i n t h e s e over  in  Considering  r e p r e s e n t a t i v e o f a c t u a l amounts of f o o d  intermittent  entrapment  located  taken  8.  these a s s o c i a t i o n s  sites  snow  e l e v a t i o n s i n the  exposed i n the  Because of p o t e n t i a l  association  bottom.  t h e s o u t h - f a c i n g s l o p e of Mount C a i n .  estimated  Mountain  associations.  was  these  and  abundance i n w i n t e r i s  overestimated  on  entrapment  f o r A m a b i l i s F i r - T w i s t e d S t a l k and  forested  probably  zone i n which  be  exposed  lower e l e v a t i o n s shrubs  much l e s s t h a n  through  that at  the  56  During generally  winter, had  logged areas. than  8).  more  Newly  elevation  food  logged, than  the  Western  Of  the  compared,  than did mainly prep. low  the  Hemlock  to  and  504  kg  s i m i l a r to ha )  logged  Fir -  associations,  component o f conifer  and  kg  ha  my  but the in the  less  estimates of  These  had  Hemlock  total  food  components.  can  food were  Rochelle  ( in  respectively.  (43  c o n i f e r s e r a i stage  s u b s t a n t i a l amounts o f more f o o d  available -  that  i n the  Salal -  most food  and  the  was  greater  did  Hemlock  Douglas-fir  three  abundant,  was food  than  Western  in  These  a v a i l a b l e deer forage The  be and  differences  Salal  available  abundance  amounts o f  and  was  Fern-  a v a i l a b l e deer forage  c o n i f e r s e r a i stage which  lichen  that  than  greater  habitats  s e r a i s t a g e had  Western  had  of  - 1  and  provided  that  shrub abundance.  forested  habitat  This  a s s o c i a t i o n . In associations  392  in  5  Salal-Douglas-fir,  associations.  at these e l e v a t i o n s .  - 1  winter.  Amabilis  and  mid-elevation  only  during  96  Hemlock,  associations  differences  ) measured  values are to  associations  plant  food  Deer  were s i m i l a r i n f o o d  six  Fir-Western  (Fig.  C e d a r , and  association.  other forested due  Red  did  Hemlock  shrub s e r a i stages  Sword F e r n - W e s t e r n  forested  more  S a l a l - Western  Hemlock-Plagiothecium  Amabilis  Salal-Western  snowpack t h a n  much l e s s i n w i n t e r  and  associations  s e r a i s t a g e s had  h e r b , f e r n , and  W e s t e r n Hemlock a s s o c i a t i o n s to  shrub  had  plant  above t h e  f i r - W e s t e r n Hemlock and  i n summer b u t  even l e s s f o o d  forested  exposed  H e r b , f e r n , and  Amabilis  associations  low  forest  the than  shrub the  57  The  amounts  forested for  plant  spring  3.3.  of  food  associations  but evaluated  during  densities  of  provided  and  and s e r a i s t a g e s were n o t d e t e r m i n e d  available to  winter  digestible  by  i n the d i f f e r e n t  (Fig. 5  dry  Rochelle  later.  D i g e s t i b l e Dry M a t t e r  d e n s i t i e s o f food summer  t o deer  q u a l i t a t i v e l y and p r e s e n t e d  D e n s i t y Of A v a i l a b l e The  available  (  and  matter  in  deer  in  habitats  6) were c o n v e r t e d t o  ( F i g . 9)  prep.)  the  using  f o r the  factors  major  forage  components. During Twisted had all  summer t h e V a c c i n i u m  Stalk,  a n d M o u n t a i n Hemlock  the highest densities the f o r e s t e d  density  plant  associations  in  Douglas-fir,  the  Amabilis  Western  cutover areas dry  Deer  winter  of the  associations  Hemlock,  were  in  -  associations i n both  to  summer  winter  Hemlock,  and  and w i n t e r . available  stages except  t h e newly  available  associations  the c o n i f e r  i n the  Hemlock,  S a l a l - W e s t e r n Hemlock, A m a b i l i s F i r  a l l s e r a i stages, except  The Sword  of  those  a n d M o u n t a i n Hemlock - C o p p e r b u s h  Salal -  have low d e n s i t i e s  the d e n s i t i e s  a l l serai  similar  Western  -Skunk C a b b a g e , A m a b i l i s F i r - W e s t e r n  Douglas-fir,  During  Western  Fern  d u r i n g summer  matter  s e r a i stage  Vaccinium  Stalk,  d i g e s t i b l e dry matter  ( F i g . 9 ) . During  Fir -  Hemlock - P l a g i o t h e c i u m  digestible  -  Fir -  associations  a v a i l a b l e i n these  a v a i l a b l e d i g e s t i b l e dry matter  logged  Copperbush  and S a l a l - Western Hemlock a s s o c i a t i o n s .  F e r n - W e s t e r n Red C e d a r ,  In  Amabilis  and may be below t h e d e n s i t i e s o f d i g e s t i b l e dry m a t t e r  available  of  -  of a v a i l a b l e  o f d i g e s t i b l e dry matter  declined  - Skunk Cabbage,  -  Salal  Twisted (Fig. 9).  serai  stage  58  Figure  9. D e n s i t y  summer  in  assessment plant Fern  of d i g e s t i b l e  habitats  sites  WP = Western  Red  above  the  AT = A m a b i l i s  Hemlock - C o p p e r b u s h  .  deer..  Fern  Cabbage,  Immature  Serai  SC =  Sword  Hemlock, Fir - Western  MC = M o u n t a i n  Stages :  S = Shrub, C = C o n i f e r .  of  Forested  AW = A m a b i l i s  Stalk,  and  Number  - Western  - D o u g l a s - f i r , SW = S a l a l  F i r - Twisted  Logged, H = Herb, F = F e r n ,  winter  histograms.  -Skunk  DW = Deer  SD = S a l a l  during  radio-tagged  VS = V a c c i a i u m Cedar,  matter  Hemlock - P l a g i o t h e c i u m ,  W e s t e r n Hemlock, Hemlock,  by  are indicated  associations : - Western  used  dry  N = Newly  60  had  low  densities  differences  of  available  i n the density  digestible  of a v a i l a b l e  between summer and w i n t e r were p r i m a r i l y herbs  and b u r i a l The  of shrubs  density  of  during the spring  dry matter.  digestible a result  dry  These matter  of the lack o f  by snow d u r i n g t h e w i n t e r .  digestible  dry  on t h e f o r e s t e d  plant  matter  available  associations  t o deer  and  serai  s t a g e s c o u l d n o t be a d e q u a t e l y  e s t i m a t e d and s o a r e n o t i n c l u d e d  in  will  this  section.  the other seasons have been The  when t h e movements and h a b i t a t  implications  will  of the  d r y matter be  relative  discussed  later.  First  use o f b l a c k - t a i l e d  are then  e v a l u a t e d with regard t o  digestible  dry matter  Seasonal  and o t h e r  data  each  seasonal  the  overlapped home  distributions  ranges of  were b a s e d  available  movements  available  factors.  from  Deer seven  radio-tagged  t h e year suggested  deer,  extensively.  that  deer  generally  to  spring,  s e a s o n a l home r a n g e s  o f each  In both  defined  home  of these by  ranges latter  different  l o c a t i o n s o b t a i n e d between c e r t a i n  on s h i f t s  movements  of  were u s e d , c o r r e s p o n d i n g  were  and  p a t t e r n s o f s e a s o n a l movements. The  over  and w i n t e r . F o r f i v e  deer  of  densities  were s e p a r a t e d . F o r two d e e r t h e s e a s o n a l  seasonal  dates  these deer  s e a s o n a l home r a n g e s  summer, deer  of  densities  a r e d e s c r i b e d . These  were c o l l e c t e d  (Table I I ) to c o n s i d e r t h e i r locations  deer  Movements Of R a d i o - t a g g e d  Sufficient  three  use o f t h e deer-  t o t h e s e a s o n a l movements o f b l a c k - t a i l e d  habitat  3.4,  be compared q u a l i t a t i v e l y t o  presented.  digestible deer  T h i s season  i n locations  of  the  deer  of  deer, areal  d a t e s . The together  61  Table  II.  range data  List  of radio-tagged  deer  and s e a s o n s  were o b t a i n e d . A l l r a d i o - t a g g e d  deer  f o r which home  were  adults.  Sector  Deer  Sex  0FL61  male  Croman  OFL68  female  Mount Cain  OFL67  female  0FL60  female  Spring  X  Summer Winter  X  Comments, Observation P e r i o d , (Number Of Locations)  May 1 9 7 5 to September 1 9 7 6 .  X  (106)  X  X  no l o c a t i o n s obtained f o r spring; k i l l e d at beginning of winter, 1 9 7 5 ; accompanied by one fawn. J u l y to November 1975.  (36)  Maquilla  X  X  X  June 1975 to June 1 9 7 6 . ( 9 8 )  Croman  X  X  X  accompanied by one fawn i n 1 9 7 6 . May 1 9 7 5 to A p r i l 1976.  0FL71 *  female  Croman-Hoomak  X  X  X  OFL58  female  Mount Cain  X  X  X  OFL62  male  Croman  X  X  X  * h o r i z o n t a l migrator  (64)  accompanied by two fawns i n 1 9 7 6 . J u l y 1 9 7 5 to September 1 9 7 6 . ( 2 1 2 ) few l o c a t i o n s obtained f o r spring; k i l l e d at beginning of winter, 1 9 7 5 . A p r i l to November 1 9 7 5 . ( 4 7 ) May 1 9 7 5 to June 1 9 7 6 . ( 1 6 5 )  63  with  t h e o c c u r r e n c e o f s e a s o n a l movements by t h e f i v e m i g r a t o r y  d e e r and n i g h t occupied  count  between March  observations. and June;  Spring  home  ranges  were  summer home r a n g e s between  June  and November; and w i n t e r home r a n g e s between November and March. These  p e r i o d s o f s e a s o n a l home r a n g e o c c u p a n c y d i f f e r e d  d e e r and a r e d i s c u s s e d  3.4.1.  Statistical  Statistical of  Parameters  evaluation  grouped were  normal  of s e a s o n a l  distribution.  estimated.  changes  deer's  according  Arithmetic  locations  i n the  to  "ring"  frequency  a  bivariate  distances  test  mean  each  (after  significantly distribution significantly  normal  of distances  deer  test  normal (Siegel  of d i f f e r e n c e s  of  the  21  different  (p  (Table  groups  of radio-tagged  seasonal  home  distribution. normality  between  deer.  range mean  were centre  Comparisons o f  were t e s t e d test  using  compares t h e  locations  distribution.  were  1966) were  and t h e  centre, t o the frequency d i s t r i b u t i o n  one-sample  Sixteen  Neft  about the a r i t h m e t i c  to bivariate  i n a bivariate  significance  approximate a  of each  ( K o w a l s k i 1 9 7 0 ) . The " r i n g "  distribution  arithmetic  in  position  parameters o f t h e s e  mean c e n t r e s  t o be d i s t r i b u t e d  seasonal d i s t r i b u t i o n s  Smirnov  The l o c a t i o n s  f o r the seasonal d i s t r i b u t i o n s  hypothesized  the  3.4,4.  Of Home Ranges  by s e a s o n and t h e d i s p e r s i o n  calculated  the  i n Section  home r a n g e c e n t r e s c a n be made i f t h e l o c a t i o n s  bivariate  The  more f u l l y  between  The  of these  Kolmogorov-  1956 : 47) was t h e n used t o t e s t between t h e two seasonal  > .05)  home  from  I I I ) . Five  (p < .05) from a b i v a r i a t e  home  a  distributions. ranges  were  bivariate ranges  not  normal differed  normal d i s t r i b u t i o n  in  6 4  Table  III. Statistical  Areal from  distributions a bivariate  test",  normal  that  are  seasonal  distribution.  the areal  o f s e a s o n a l home  ranges.  significantly different  normal d i s t r i b u t i o n  incorporating  compare t h e  characteristics  are  underlined.  Kolmogorov-Smirnov t e s t , distributions  with  the  (p<.05)  The was  "ring used  to  bivariate  Standard D i s t a n c e D e v i a t i o n ( m ) spring OFL61  376.6 (34)  OFL68  -  summer 1 0 0 1 . 6 (39)  (10)  491.0  (38)  (10)  OFL60  311.6  679.2  (20)  •  "  (33)  649.9  495.7  "  534.7  (26)  2 5 0 . 3 (50) (15)  winter  395.1  OFL67  w i n t e r range  a u x i l i a r y w i n t e r range  398.2  (12)  412.1  (17)  0FL71*  157.4  (19)  490.5 (167)  262.2  (26)  0FL58  171.6  (3)  9 2 4 . 5 (29)  370.6  (15)  0FL62  484.2 (50)  508.1  455.2  (69)  (46)  + number o f l o c a t i o n s are e n c l o s e d i n b r a c k e t s * horizontal  migrator  Skewnees / K u r t o s i s  +  spring .18 /  summer .38  .12 /  .50  .44 /  .05  winter  .87 / -.10  .28 /  .34  .44 /  .30  .34 /  .71  .25 /  .51  .26 /  .42  1.08 / -.46 .24 / - . 0 7  ;  /  .29  -.19  .20 /  .82  .12 /  .57  / -.50  .24 /  .81  .47 /  -.25  .31 /  .03  .19 /  -.26  .73 /  1.36  .17  .19 /  .71  66  that  they  were  a l l leptokurtic  c e n t r e and a t t h e t a i l s slightly of  skewed  than  (slightly  (more l o c a t i o n s  in  a  normal  asymmetrical).  t h e summer home r a n g e o f OFL67, t h e  observed  distribution  distribution,  and  was l i k e l y  the  caused  because  of  locations  poor  on o n l y  precise  location  several  days's  locations a small  radio  reception,  gave  that  difference  in  a  remote  normal  area  and,  t o get accurate OFL67,  when  on t h e b a s i s o f  resulted  i n several  and, when c o u p l e d  appearance  the  i n l o c a t i o n . The  was e s t i m a t e d estimation  the  between  bivariate  I was a b l e  a t t h e same p o s i t i o n  size,  I n one i n s t a n c e ,  The l o c a t i o n o f  was n o t p o s s i b l e ,  observations. This  sample  and  by i n a c c u r a c i e s  a few o c c a s i o n s .  occurring  distribution)  hypothesized  summer home r a n g e o f 0FL67 was l o c a t e d  n e a r t h e mean  of  a  with  leptokurtic  distribution. Of  the  remaining  significantly ranges,  (p  OF158  < .05)  four from  home  ranges  bivariate  summer a n d OFL62 s p r i n g ,  probability  level  of  0.01.  In  differences  from  the  bivariate  significant,  were  not  substantial  that  normality  differed two  home  were n o t d i f f e r e n t a t a  a l l four normal with  home  ranges,  the  d i s t r i b u t i o n although kurtosis  reaching  a  maximum o f 0.82 and skewness a maximum o f 0.26.  3.4.2.  Home Range  Home the  range  Size s i z e was c a l c u l a t e d  minimum home r a n g e method f i r s t  (1938) convex during  (Table  IV).  polygon  using  modified  p r o p o s e d by D a l k e  versions of and  Sime  The 10055 home r a n g e i s t h e a r e a e n c l o s e d  by a  surrounding t h e outermost  a p a r t i c u l a r season.  This  i s  the  locations most  o f t h e deer  commonly  used  67  Table  IV-  eliminate  Size  of  seasonal  home ranges. The 90% home ranges  i n f l a t e d home range s i z e s caused by extreme  The 50% home range i s the area of i n t e n s e deer use.  locations.  S i z e o f Home Range (ha) Number o f locations  100  90  %  SP  SM  WT  SP  SM  WT  SP  0FL61  34  39  33  89.3  4o6.6  186.7  38.6  OFL68  -  26  10  66.9  49.7  0FL67  50  10  38  63.2  79.3  0FL60  15  20  "  27.0  w i n t e r range  12  " aux. winter range  17  0FL71*  19  167  _  69  146.1  56.0  migrator  147.5 104.5  165.7  13.3  21.3  16.8  14.9  88.4  65.3  30.7  10.6  9.0  .6  4.4  -  4.6  28.3  5 . 5  8.2  16.0  46  34.0  4.2  WT  33-9  152.0  50  84.7  SM  59.5  .3  0FL-62  274.1  SP  2.6  36.6  15  WT  20.3  185.6  29  SM  %  49.5  6.7  3  mean  138.I  26  0FL58  * horizontal  66.8  50  %  72.7  5.5  36.4  12.5  20.0  12.0  52.6  88.9  83.O  26.6  79.8  40.6  .5  13.5  2.4  4.1  2.7  6.8  29.1  26.5  4.1  19.1  7.4  69  estimate  o f home r a n g e s i z e .  sensitive  t o extreme o r abnormal l o c a t i o n s  order  to circumvent  range  . This estimate  polygon  this  sensitivity,  the  e l i m i n a t e s the i n f l a t i o n  closest  i n area  when t h e 100% method i s u s e d . is  polygon in  the  50%  of  the  I calculated  the core  90%  of  caused  A third  home r a n g e  enclosing the closest  identifying  o f t h e 100% home r a n g e i s  o f home r a n g e i s t h e a r e a  surrounding  range  The s i z e  In  t h e 90% home  enclosed  the  by  animal.  by  locations  extreme  a  and  locations  e x p r e s s i o n o f s i z e o f home  . This estimate  i s the area  of a  50% o f t h e l o c a t i o n s and i s u s e f u l  of deer  use w i t h i n t h e 100 o r  90%  home  i n Figures  10 t o  ranges. The 16.  90%  home r a n g e s  The 50 and 100% home r a n g e s  locations  of  the  areal  expression  Since  the area  r a d i u s then  deviation  z  are  of v a r i a b i l i t y  in a  deer*s  Neft  1966)  the  regression  was  was s i g n i f i c a n t l y  0.61  hypothesized  100%  100%  and  the  home  range  i f there  of i t s  distance  2.52 ±  size  was  slope  17).  of the  (p < . 0 5 ) .  0.97,  included  limits.  a function of standard  f o r homogeneity of v a r i a n c e  were s e a s o n a l  t h e 100%  (Fig.  from z e r o  v a l u e o f 2 i n i t s 95% c o n f i d e n c e  deviation, a test  home r a n g e s  variable  different  the  determine  position.  includes a l l locations,  exponent o f t h e r e g r e s s i o n e q u a t i o n ,  to  average  be p r o p o r t i o n a l t o t h e s q u a r e  The  distance  although  i s p r o p o r t i o n a l t o t h e square  was used a s t h e d e p e n d e n t  regression line  Since  delineated,  distance d e v i a t i o n . Since the standard  (after  of  not  a r e i n d i c a t e d . S i z e o f home r a n g e i s an  home r a n g e s i z e s h o u l d  home r a n g e a r e a r  deer  of a c i r c l e  of the standard  The  ( T a b l e IV) a r e shown  was  used  differences i n the size of  ( T a b l e s IV a n d V ) . A s i d e  from t h r e e  exceptions,  70  Figure  10. Seasonal  90% hose  ranges  of  altitudinal  migrator  OFL6 1. Shaded areas are logged, unshaded areas are unlogged. 90% s p r i n g  ^  home range  o  spring  location  90% summer home range  •  summer l o c a t i o n  90% winter home range  •  winter  location  72  Figure  11-  Seasonal  90%  hoae  ranges o f a l t i t u d i n a l  migrator  OFL68. Shaded areas a r e logged, unshaded a r e a s a r e unlogged. ===== 90S summer home range  •  summer l o c a t i o n  11 90% winter home range  •  winter  location  74  Figure  1 2 . Seasonal 9 0 S  OFL67. s h a d e d  ranges  of  altitudiaal  aigrator  a r e a s a r e l o g g e d , unshaded a r e a s a r e unlogged.  {jjU|j}|{ 9 0 * s p r i n g ^^^^  hoae  hoae r a n g e  o  spring  location  9 0 * summer home r a n g e  •  suamer  location  •  winter  location  ^90%  w i n t e r home r a n g e  75  76  Figure  13.  Seasonal  90%  hoae  ranges of a l t i t u d i a a l  OFL60. Shaded areas are logged, unshaded a r e a s are jjjjjijjjj 90% s p r i n g •  migrator  unlogged.  hoae range  o  spring  90% suamer hoae range  •  summer l o c a t i o n  •  winter  ^90%  winter hone range  location  location  78  F i g u r e 14. Seasonal  90%  hose  ranges  of  horizontal  aigrator  0FL71. Shaded areas a r e logged, unshaded areas a r e unlogged. Illlllllll  9 0 5 4  spring  hone range  o  zEEEEEz 90* suamer home range H  90% winter hoae range  spring •  •  location  summer l o c a t i o n winter l o c a t i o n  79  80  Figure  15.  Seasonal 90% home ranges  o f r e s i d e n t 0FL58. The 90%  s p r i n g home range i s not shown nut i t o v e r l a p s e x t e n s i v e l y the  90%  summer home range. Two extreme l o c a t i o n s i n summer a r e  i n d i c a t e d with mean centres ^  with  •  ; o t h e r l o c a t i o n s a r e not shown. A r i t h m e t i c  of s e a s o n a l home ranges a r e i n d i c a t e d by:  spring,  summer,  ^  winter.  logged, unshaded areas a r e unlogged.  90% summer home range 90% winter home range  Shaded  areas  are  82  Figure  16.  Seasonal  A r i t h m e t i c mean centres  90%  borne  ranges  of s e a s o n a l hone  of ranges  resident  OFL62.  are i n d i c a t e d  by: ^  spring,  l o g g e d , unshaded  summer,  ^  winter.  areas a r e unlogged. I l l l l l l l l l 90% s p r i n g home range 90% summer home range i f 90% winter home range  Shaded  areas a r e  84  Figure size  17. R e l a t i o n s h i p of  100%  home  r = . 6 1 , n=21; where 2  distance  deviation.  between standard range.  H  i s home  distance  deviation a  H = .000014 s - 5 2 ± . 9 7 , p<.0 2  range  s i z e and S i s standa  86  T a b l e V. (Sokal  Differences in size and  deviations.  Rohlf  1969  of  :186)  100% to  home r a n g e s u s i n g compare  standard  an  F-test  distance  d.f.  f  0FL61  38,33  7.07  1.69  ^ . 0 5  OFL68  -  -  -  0FL6?  9,49  3.92  0FL60  19,14  4.75  —  +  166,18  OFL58  28,2  OFL62  45,49  +  f  32,33  2.01  -  -  f  38,32  3.51  1.79  ^.05  -  9,25  2.70  2.28  2  2.12  5 . 0 5  9,37  1.02  2.12  >.05  37,49  3.84  1.74  2.46  s .05 19,11  2.91  2.65  05  11,14  1.63  2.57  > .05  19,16  2.71  2.35  s.05  16,14  1.75  2.46  ^.05  16,11  1.07  2.72  >.05  a u x i l i a r y w i n t e r range  9.71  1.66  29.02  19.50  1.10  1.56  f = f calculated c  * horizontal  d.f.  d.f.  P  c  , '  migrator  P  t  .05  between winter ranges  t  P  1.84  •s.05  f  c  -  .05  166,25  3.50  1.61  ^.05  25,18  2.77  2.15  •s.05  .05  28,14  6.22  2.35  s.05  14,2  4.66  19.40  >.05  >.05  45,68  1.24  1.59  >.05  49,68  1.13  1.56  >.05  5 . 0 5  2  f  c  —  0FL71*  Winter - S p r i n g  Summer - Winter  S p r i n g - Summer  f.= f t a b l e t  88  summer home r a n g e s  were l a r g e r  than  either  winter  o r s p r i n g home  ranges. Distance (p > .05) This  variances  between  similarity  small  the  of  not  (n = 3) used  her  spring  probably  inaccuracies  in  home  range.  relocation  o f OFL67 may be  the  throughout largely  the year.  a  result  0FL62  which  These s i m i l a r i t i e s of  the  more  stayed  and  home r a n g e  distance i n size  a  result  range  size  t h e same  size  i n home r a n g e s i z e  remote  locations  C o m p a r i s o n o f h i s 90 o r 50% home r a n g e s i z e s spring  the  a s d e s c r i b e d i n S e c t i o n 3.1.1.  were  of  from  The s i m i l a r i t y  e x c e p t i o n s t o g e n e r a l t r e n d s i n s e a s o n a l home ranges  o f OFL58.  results  Final  home  different  to estimate the standard  between summer a n d w i n t e r home r a n g e s of  significantly  s p r i n g and w i n t e r home r a n g e s  i n home r a n g e s i z e  sample s i z e  deviation  were  of  indicated  were OFL62.  that  was t h e s m a l l e s t o f h i s s e a s o n a l home  his  ranges,  s o was c o n s i s t e n t w i t h t h e g e n e r a l t r e n d s p r e v i o u s l y s t a t e d .  Although winter  t h e summer home range o f home  observed  range  i n other  The  the  difference  in  spring,  90%  home r a n g e s ranges,  7  spring,  half  averaging During  the  was n o t a s g r e a t a s t h a t  deer  were between 47 and 55% t h e s i z e  and  than  was  56.0  165.7 ha i n summer, and 72.7 h a i n w i n t e r . Mean  and  illustrated  12% t h e s i z e  of a radio-tagged  4.1  larger  f o r t h e seven  the  a b o u t t h e a r i t h m e t i c mean c e n t r e . between  was  deer.  mean 100% home r a n g e s i z e  ha  home  ,  OFL62  ha.  In  of  mean  c o n c e n t r a t i o n of l o c a t i o n s  Mean  50%  home  ranges  o f mean 100% home r a n g e s . dee^s  summer,  100%  locations  were  were  Thus, i n in  areas  t h e a r e a i n c r e a s e d t o 19.1 h a .  t h e w i n t e r , t h e . c o r e . o f deer  use a s d e l i n e a t e d by t h e 50%  89  home r a n g e d r o p p e d  in size  These  of d i f f e r e n t  comparisons  consistent  with  were s i m i l a r  3.4.3.  the  to  seasonal  results  Of  home  percentage  ranges,  can  that  arithmetic  mean c e n t r e s  they  the  the  from  (Neft are  same.  the in  derivation  also  be r o b u s t With  In  means  of  o u t s i d e o f the from the  a  The  with  regards  table  can  was  from  were  be  used  i n t e r v a l between t h e t h e u p p e r and ( S o k a l and  a p p l i e d t o the  by  normal  different, that  the  to normality  (1966) i s  of  similar  normality.  v a r i a n c e s are to test  for  calculated  critical lower Rohlf  t -test v  Neft  shown t h a t  noted  regards  the  T h u s , I assumed i t s h o u l d  bivariate  s a m p l e s . However, t h e  using  (1973)  test  that  bivariate  significantly  with  t-test.  a  the  different  and  S e c t i o n 3.1.1., i t was  d a t a , i f the  still  d e g r e e s of f r e e d o m  constraint  to  with  two  test  normal  Gilbert  Student's  i n o r d e r to  bivariate  proposed  r  comprising  c o n d i t i o n s of the  robust  t -test  to the  t-test  ways.  is fairly  univariate  Student's the  those  i n minor  sample d a t a .  where home r a n g e s  1966). The  that  and  t-test  were  populations  distribution  Student's  home r a n g e s i z e s  compared  different  only  ha.  locations,  most home r a n g e s were n o t  differed  7.4  distributions.  be  come  distributions  are  covered  o f ' S e c t i o n 3.1.1.  data,  hypothesis  variances  average,  Home Ranges  univariate  are t h a t the  on  b i v a r i a t e normal  Position  as w i t h  and,  of  t  sample  different,  the  differences  in  t value  must  be  values  derived  s i z e s to  1969:375). Neft  (1966).  This  obtain same  90  For  each  radio-tagged  mean c e n t r e s o f t h e different  seasonal  in  summer and  winter  difference  home  home  due  to  ranges  the  were  standard  distance  be  detected.  sample  size  . A second  (n  factor  This v a r i a b i l i t y  was  ranges.  resulted  =3)  was  sighted fawn.  After  The  high  OFL58  moved  standard  lost  these  movements  shot  dead  t h e e x t r e m e movements, OFL58  between  seasonal  o f home r a n g e s were used  as  migratory  Migratory  only  OFL58.  I  one  Within to her  u n t i l her signal season.  between t h e a r i t h m e t i c mean c e n t r e s o f w i n t e r and  positions  VI).  by  returned  there  d i s t a n c e s between  home r a n g e s a n d w i n t e r o r s p r i n g home r a n g e s Distances  extreme  extreme l o c a t i o n s ,  400 m f r o m  summer r a n g e a n d r e m a i n e d  that  ( F i g . 1 5 ) . OFL58  accompanied  home r a n g e s were much l e s s t h a n  either  of  distance deviation i n  on t h e s e c o n d d a y o f t h e a n t l e r l e s s h u n t i n g  Distances spring  to  i n the cutover,  A n o t h e r fawn was f o u n d  elevation  was  high  may a l s o h a v e c o n t r i b u t e d t o t h e l a c k  f r o m d i s t u r b a n c e by h u n t e r s  her standing  a few d a y s a f t e r low  used t o  the  a c c o m p a n i e d by two fawns a few days p r e v i o u s t o t h e  movements.  of  between t h e a r i t h m e t i c mean c e n t r e s o f h e r summer a n d  home  have  Lack  d e v i a t i o n o f OFL58's summer l o c a t i o n s ( T a b l e  OFL58's summer l o c a t i o n s was due t o two extreme may  significantly  10 t o 1 6 ) . F o r OFL58, no  could  small  h e r s p r i n g home r a n g e  winter  ranges  between h e r s p r i n g and summer home r a n g e s  calculate  difference  f o r OFL58, a r i t h m e t i c  t h e p o s i t i o n o f s p r i n g and summer home r a n g e s o r  i n position  likely  III).  except  (p < .01) ( T a b l e VI a n d F i g s .  difference  was  deer,  deer  home  had s e p a r a t e d  (Table V I ) .  ranges and t h e r e l a t i v e  to classify  o r r e s i d e n t deer  summer  (Figs.  radio-tagged  10 t o 16  and  summer and w i n t e r home  deer Table  ranges.  91  Table  VI.  Distance  between  arithmetic  r a n g e s and t e s t s o f d i f f e r e n c e s i n t h e i r  mean  centres  positions.  o f home  distance (km)  t  r  Winter - S p r i n g  Summer - Winter  S p r i n g - Summer P  distance (km)  distance  P  OFL6I  4.54  24.62  ^.01  3.64  18.46  •=.01  .91  OFL-68  -  -  -  4.30  23.42  s.01  -  1.44  8.07  s.01  range  2.02  9.07  a u x i l i a r y w i n t e r range  4.05  20.87  OFL67  2.23  20.76  S.01  OFL60  2.45  12.62  5.01  winter If  7.99  i.01  -  -  .79  9.74  S.01  S.01  .44  3.08  2.01  £.01  2.79  20.68  -  2.93  18.45  < .01  between winter ranges  0FL71*  6.16  54.15  P  (km)  .01  ^.01  5.89  59.59  ^.01  .28  4.06  s  .01  0FL58  .40  .72  >.05  .09  .36  >.05  .31  2.68  ^.01  0FL62  .34  3.33  s.01  .28  3.07  5.01  .62  7.10  s.01  h o r i z o n t a l migrator ro  93  while  their  s p r i n g and w i n t e r  or overlapped. range  Resident  centres,  basically  3.4.4.  although  t h e same t h r o u g h o u t  dates  o f home r a n g e s  the  locations.  tagged home  ranges  deer  p e r i o d s chosen, (Section  therefore  occupied  in  migratory  and w i n t e r  the  study  periods  distances  spring spring  occupancy  of  arbitrary.  resident  However,  activity  dates  i n the, c o r e o f radiodeer's  during  were  occurred  i n the dates  b u t g e n e r a l l y s p r i n g home  moved  home r a n g e s  area,  during  movement o f d e e r  were  on s h i f t s  of  changes i n  of migration of other  centres of  differences  m i g r a t i o n s were p r e s e n t e d the  dates  ( T a b l e VI)  between March a n d J u n e , summer home r a n g e s  November,  Thus,  both  the  the  identified  3.4.3.).  s e a s o n a l home r a n g e s ,  and  remained  a r e shown i n  by d i s t i n c t  I n t h e two r e s i d e n t d e e r  different  Individual of  ( T a b l e VI)  were i d e n t i f i e d  The e x a c t d a t e s o f were  home  Home Ranges  l o c a t i o n s and on d a t e s  deer.  i n their  home r a n g e b o u n d a r i e s  o f home r a n g e o c c u p a n c y were b a s e d deer's  separated  t h e year.  migratory  occupancy  the  shifts  o f o c c u p a n c y i n s e a s o n a l home r a n g e s  18. I n t h e f i v e  deer's  were e i t h e r  made s e a s o n a l  their  Occupancy Of Seasonal  The Figure  deer  home r a n g e s  year.  ranges  deer  The  were  between J u n e  between November  black-tailed  the  of occupancy  and March.  exhibited three  distances  of  these  i n S e c t i o n 3.4.3. S p r i n g m i g r a t i o n was  between w i n t e r  and s p r i n g home r a n g e s .  were s m a l l , b e i n g u s u a l l y  0.9 km,  but  a s g r e a t a s 2.8 km. Summer m i g r a t i o n was t h e movement  from  t o summer home r a n g e s , migration,  with  less  and was u s u a l l y  distances  extending  than  The  greater  than  t o 6.1 km.  the  Winter  94  Figure  18.  SP = s p r i n g , Short  Periods  occupancy  SM = summer,  vertical  Consecutive  of  lines  are  of  WT = w i n t e r , dates  l o c a t i o n s on a s e a s o n a l  when  seasonal  home  AW = a u x i l i a r y deer  were  range a r e j o i n e d .  ranges. winter. located.  95  HIH I  SP-j SM-  Hi  f lIII I 1 —I  1  1  OFL61  4  i in iii mu i i m—H4-  WT  1  II IE  1  1  II—hH—III  1  1  1  M  MU  III "  1  1  '  1  I  1  I  SP-,  1  1  OFL68  SM-  (  I I  I I 11  1  1  I H  WT -i  cr  o cr  o  1  1  1  1  -i  r-  SP-i  • I IB  SM-  II  HI 'ai »i  1  1  OFL67  •H MM*  WT  III I 11 I—l-IIHIBll -I  SP-i  I  1  1  l(  l  r-  1  i  1  1  '  1  i  i—6-rt  SM-  VII  I n—i  H—i—«H(  WT  OFL60  III I I I I — r  AW-  Vi i .  -i  ,  1  1  1  1  >  r  1  ill  1  1  1  '  1  i  i ~  SP-  '  1  OFL71  SM-  i l a II  m i nam II 11—III lain 111  i — — M  WT T  -i  r  1  1  1  -i  r  —r  1  1  1  1  SP-j  -z.  Q  if)  UJ cr  r  —  OFL58  SM-  llll I I I I III I  1—HHI  WT-I  1  1  1  -I  r  1  1  1  SPi  1  1—  1  '  f  011II W" I  SM-  OFL62  Mil I II I I I — H - t — H  WT  Mn i -i  A M J 1975  1  J  r  II  ~i  A S O N D  i — i in i wu 1  J  1  III  mm-  1  1  1  F M A M  1976  1  —  i  1  1  J J A S  i  i  96  migration The  was t h e movement f r o m  distances  slightly  moved  s h o r t e r than  One  deer,  in  the  those  near  i t s s p r i n g home r a n g e f o r the  mouth  first  Croman  and  winter  Valley.  then  home r a n g e  Another deer,  again  first  part of winter before  The ranges  movement  may o c c u r  period.  In  of  a  s e v e r a l deer, short  home  moved i n m i d -  sidehill  at the  with  two  consequent  w i n t e r home r a n g e . February  on  to her a u x i l i a r y  have a  locations  the  winter  mid-winter  migration  i n c l u d e d i n t h e summer observed  during the  0FL71 moved t o h e r w i n t e r  home r a n g e  l o c a t i o n s were  an a u x i l i a r y  more t h a n  c h a r a c t e r i z e d by  early  ( F i g . 14) were m a i n l y  movement t o i d e n t i f y  this  the spring migration.  t h e Hoomak s e c t o r . I n s u f f i c i e n t  this  deer  there approximately  weather  in  0 F L 7 1 , may a l s o  0FL71  the  t o her e a r l y  until  range  was  the  This  i n t h e Croman s e c t o r  and t h e n  r a n g e on  in  t o OFL60's. The extreme of  migration.  , she returned  home  in  usually  She r e m a i n e d  returned  r a n g e and r e m a i n e d t h e r e  similar  were  the s i d e h i l l  winter  When snow began t o a c c u m u l a t e early  migration  ( F i g . 1 3 ) . OFL60 o c c u p i e d  weeks u n t i l a v e r y m i l d s p e l l snowmelt,  ranges.  p a r t of the winter  t o her a u x i l i a r y of  home  OFL60, e x h i b i t e d a f o u r t h  a w i n t e r home r a n g e a l o n g  January  winter  winter  moved d u r i n g t h e summer m i g r a t i o n .  had  range  summer t o  deer  taken  during  w i n t e r home r a n g e .  between two o f i t s s e a s o n a l  once d u r i n g  a  particular  migratory  movement between s e a s o n a l home duration  visits  before  home  ranges  permanent  occupancy  o f t h e new s e a s o n a l home r a n g e o c c u r r e d  example,  0FL61 was l o c a t e d on h i s s p r i n g home range on J u n e 21,  1976. Twelve  On J u n e hours  23 he was l o c a t e d later,  on  h i s summer  (Fig. 18).  home  For  range  he was back on h i s s p r i n g home r a n g e  where  97  he  remained  relocated night  he  there  until on  had  OFL68,  until  July  these  of  winter  winter  home r a n g e s  This  suggests  that  a direct  , hut  and  was  6 he  corresponded  home  ranges. occurred  deer  deer  3.6  in less  km  by  lack  with  m i g r a t i o n was  of  the  occupancy 0FL61 at  the  although  the  visits  was  well  movements between summer  and  than  12 h o u r s . 4.3  km  by  every  summer  along  the  and  the  The  distances  OFL68. 12 h o u r s winter  migratory  i n l e s s than  Even  routes  12 h o u r s  migrations  o f 0FL61 and  OFL68  blocks of  are  reserved  for  i n h a b i t e d an  that  was  was  extensively  connection route  migration c o r r i d o r s .  t o low  between  between f i v e anqustifolium  logged  and  did  not  e l e v a t i o n s . Cutover  the  summer and  winter  s i x y e a r s o l d and  . OFL68 was  0FL61  forest  w h i l e OFL68 i n h a b i t e d an a r e a  and  located  and  ranges.  consideration since substantial  unlogged,  few home  merits further  virtually  and  to  performed  winter  to h i s  snow l i n e  movement between s e a s o n a l  Comparison  In  ( F i g . 19).  0FL61 and  locations  located  snowfalls  were sometimes m o n i t o r e d  of  that  November.  moved below  Some  was  by  returned  migration.  were o b t a i n e d between t h e i r  ranges.  was  winter  summer home r a n g e s  were about  locations  to  OFL61  r  day  snow d e p o s i t i o n d u r i n g i n i t i a l  their  these  range  29  p r i o r t o permanent s e a s o n a l  w i n t e r home r a n g e s ,  travelled  home  Then on J u l y  d u r i n g the  movements  limit  though  4.  duration v i s i t s observed  above t h e  L a t e r on June  remained t h e r e through  e l e v a t i o n s of t h e i r lower  27.  moved t o h i s s p r i n g  summer r a n g e and  were a l s o  June  h i s summer home r a n g e d u r i n g t h e  daily  Short  at least  have  along  home r a n g e s  only  that  a direct forested  areas  vegetated  area  mainly  once  the  direct  o f OFL68 were by  outside  E£ilobium of  its  98  Figure  19.  migratory their  deer,  winter  ranges.  between summer and w i n t e r  0FL68 and 0FL61, s h o w i n g t h e migrations  to  snowfalls  on  station,  while  the  t o t h e 1098 m snow-depth  home ranges, by  correspondence" their  The summer home r a n g e of OFL68 was c l o s e  snow-depth close  Movements  to  of  summer home the  9:15 m  summer home r a n g e o f 0FL61- was  station.  :  99  0  i 16  1  i 18  O C T  1  i 20  1  i * i * i * i 22  24  26  1  28  i  1  30  i  i  1  1  1  i  3  N O V  1  5  i  1  i 7  I  1  9  1  11  I I 1  13  15  100  summer  or  winter  c u t o v e r about ranges,  i t appears  to  reach  the  cutover  range  her  migratory  completely  in  by  located,  about  behaved s i m i l a r l y logged  by  area  did  deer  bottom  o f t h e study  other  reflected  c l i m a t e i n the Davie Hoomak  i t s spring  northern late  the OFL71  home of the  suggest  that  i n h a b i t e d an  i n h a b i t e d an a l m o s t  spring  home the  occupied  i t s spring  the l o c a l  home  longer  persisting  range  i t s spring  of  than  slightly  later  snowpack  i s more s e v e r e  they  deer t o spring  since the than  in  i n t h e Croman s e c t o r . OFL60,  half  Croman  Valley,  by m i d - A p r i l , l a t e r  than d i d  home  way  range  up  in  mid-May.  was c o l d e r and snow p e r s i s t e d  the  other  than  radio-tagged  This  bottom  climate  i n t h e Hoomak s e c t o r  range.  Kiver valley  home  of the l o c a l  budded a n d f l u s h e d e a r l i e r  p o r t i o n of the v a l l e y  occupancy  of  severity  o f March. T h e r e ,  a r e a . L e s s snow f e l l  c l i m a t e i n Croman V a l l e y  The  deer  occupancy  reflect  s e c t o r but milder than  OFL67. 0FL61 o c c u p i e d  the  summer  even t h o u g h one d e e r  whose s p r i n g home r a n g e i s a b o u t occupied  ( F i g . 19)  s e c t o r s . OFL67 was t h e n e x t  i t s spring  migration  the  the  o f t h e Hoomak s e c t o r i s m i l d e r  w i n t e r , and s h r u b s  occupy  order  15 cm.of snow. C o m p a r i s o n  while t h e other  a t the beginning  the valley  in  although  o f permanent s e a s o n a l  migratory  range  during  in  f o r e s t e d area.  date  portions  the cutover  home  w i n t e r home r a n g e . No snow was on t h e g r o u n d i n  c l i m a t e s on t h e s p r i n g home r a n g e s . home  was i n t h e  r o u t e between t h e s e  t h a t she moved t h r o u g h  p a t t e r n s o f OFL68 and OFL61  extensively  ranges  Since t h i s l o c a t i o n  way a l o n g t h e d i r e c t  covered  deer  The  ranges.  where she was  was  these  half  home  than  spring  i n the southern home  range  The  longer i n portion.  by 0FL61 was  101  consistent  with t h i s  variation  D i f f e r e n c e s i n t h e date similar  t o although  occupancy  of  not as  spring  home r a n g e s  reflected  snowfalls.  In  at  earlier  as  ranges.  observed  and s e v e r i t y  than  deer  with  moved  10 t o 13 and  due t o t h e a b r u p t  to  of  their  18).  those  nature  The  observed  the  first than  winter  home  a t m i d d l e and  time  differences  f o r the occupancy  These s h o r t e r  differences  o f s n o w f a l l s compared t o t h e  l o n g e r p e r i o d s i n v o l v e d i n snowmelt and t h e  gradual  uncovering  p h e n o l o g i c a l d e v e l o p m e n t o f f o r a g e p l a n t s d u r i n g s p r i n g and  early  summer.  3.4.5.  Topographical  The 3.4.3  Features  Of S e a s o n a l  Home Ranges  two p a t t e r n s o f s e a s o n a l movements d e s c r i b e d i n S e c t i o n  were t h o s e  centres  of  of difference  o f m i g r a t i o n and o f m i n o r  activity.  seasonal  For  between c e n t r e s o f a c t i v i t y  evident, types  migratory altitudinal  were  migrators  deer,  (Table  with  elevation  than  in  and t h e 10 t o 1 6 ) .  movements were  migrators.  These  by t h e m a g n i t u d e o f c h a n g e s i n e l e v a t i o n  (Table V I I ) .  spring locations, either  VI)  (Figs.  two t y p e s o f s e a s o n a l  m i g r a t o r s and h o r i z o n t a l  identified  made d u r i n g m i g r a t i o n  shifts  T h e s e p a t t e r n s were d e f i n e d by t h e d e g r e e  d e g r e e o f o v e r l a p o f t h e home r a n g e b o u n d a r i e s  in  f o r the  of the winter  summer home r a n g e s  t h e s p r i n g and summer home r a n g e s .  and  were  e l e v a t i o n s where snow o c c u r r e d e a r l i e r  were much s h o r t e r t h a n  are l i k e l y  those  The o c c u p a n c y  the occurrence  e l e v a t i o n s (Figs.  involved of  great  home  climate.  o f summer home r a n g e o c c u p a n c y  low e l e v a t i o n s ( F i g . 2 ) , d e e r  ranges low  high  in local  their  For  the  three  s p r i n g home r a n g e s  summer o r w i n t e r home  altitudinal were  lower  ranges.  1 0 2  Table  VII. Elevations at the arithmetic  ranges  showing  altitudinal  horizontal  (OFL71) m i g r a t o r y  can  altitudinal  make  home r a n g e s o v e r l a p  (0FL61, patterns.  changes  in their  mean OFL68,  centres  of  home  OFL67, OFL60) and  Although  resident  deer  home range c e n t r e s  their  and s o a r e n o t a l t i t u d i n a l  migrators.  Change I n E l e v a t i o n ( m )  Elevation ( m ) spring  summer  winter  30k  1094  4o8  OFL68  -  878  437  OFL67  358  686  488  OFL60  480  725  0FL61  II  +790  summer t o winter  -686  winter to spring  -104  -441 +328  -198  -130  +245  •  _ w i n t e r range  559  -166  -79  w i n t e r range  670  -55  -190  11  auxiliary  0FL71*  282  286  OFL58  570  555  OFL62  288  431  * horizontal  spring to summer  migrator  +6  +4  -10  681  -15  +126  -111  628  +143  +197  -340  276  104  Summer home r a n g e s in  elevation  (Table lower  both  i n elevation  than  winter  late  centres  winter  home  of winter  valley  sides  at  move  to  down  range  was  elevations.  floor,  spring.  T h i s p a t t e r n corresponded  depth  and  other  over  there  was s h a l l o w  ranges  were a r e a s  lasting duration  than  Although  migrator OFL71  snow  deeper  Croman  pattern  ranges  to the valley  spring  on  the  sidesf o r  pattern  ( F i g . 2 ) . Summer  snow c o v e r . snow  depth  snow where  Spring  cover  of  ranges  were a t e l e v a t i o n s  but of shallower  of  e l e v a t i o n s and  t o the a l t i t u d i n a l  centres of the seasonal  home  longer  and s h o r t e r  home r a n g e s o f  0FL71, were a l l w i t h i n 10 m o f one not  make  m i g r a t i o n , she d i d accomplish moving f r o m  were  ranges.  the  did  aspect  migrators  s t a y on t h e v a l l e y  where snow was d e e p e r a n d  at  since  was deep snow a n d c o n t i n u o u s  winter  ranges  the  The  general  t o spend  55 m  aspect.  slopes adjacent  snow a n d i n t e r m i t t e n t  on summer  Elevations horizontal  while  on w i n t e r  than  were d i f f e r e n t  northern  The  d u r a t i o n o f snow c o v e r  winter,  ranges  However,  summer a t h i g h  deer  were a t e l e v a t i o n s where t h e r e cover  a  move down f u r t h e r  while  range.  of a l l altitudinal  valley  snow  home  on  i s t h a t they  then  home  home r a n g e was o n l y  home r a n g e s  w i n t e r on s i d e h i l l  Some d e e r  spring  higher  range o f OFL60 was on a s o u t h e r n  low  migrators  m i g r a t o r s were  and  winter  summer  home r a n g e s  altitudinal  floor.  the  and v e g e t a t i o n o f these  w h i l e h e r summer home  on  their  V I I ) . I n OFL60, t h e l a t e  climate the  than  of the a l t i t u d i n a l  Valley,  an a  elevational  change  in  where t e m p e r a t u r e s  change  local  another. during  climate  by  a r e c o l d e r and t h e  and more p e r s i s t e n t , t o t h e Hoomak s e c t o r where t h e  c l i m a t e was m i l d e r .  105  The  r e s i d e n t deer,  patterns  of  changes  characteristic three  elevation  of a l t i t u d i n a l  exhibited  i n elevation  a different  the v a l l e y valley  included  bottom  bottom  in  the  spring  home  centred  higher  The  in  not  exhibit  the  home r a n g e s e p a r a t i o n  Although  based  on  only  than  summer  used  during  provide  occur  foraging  than  differed  more r e s i d e n t d e e r shift  their  home  143  spring,  was  also  m  higher  than  the  because  from  the  winter  cutover  altitudinal home r a n g e s  migratory were  higher  summer home r a n g e s .  areas  f o r deer  were t a g g e d  was  h i s summer home r a n g e .  the centres of their  habitat  the  some o f  The w i n t e r home r a n g e o f OFL62  i n t h a t t h e c e n t r e s of t h e i r  p a t t e r n may  of  , t h e c e n t r e o f t h e summer  sidehill,  on t h e s i d e h i l l  than  range  o f 288 m. A l t h o u g h  home r a n g e  centre.  r e s i d e n t deer  elevation  home  h e r summer home r a n g e . OFL62  a t an e l e v a t i o n area  range  . The w i n t e r  p a t t e r n . H i s s p r i n g home r a n g e was l o c a t e d  home r a n g e was on t h e l o w e r  deer  and  migrators.  o f h e r summer home r a n g e  OFL58 was h i g h e r  this  in  did  l o c a t i o n s t h e s p r i n g home r a n g e o f 0FL58 was c l o s e t o t h e  elevation  in  OFL58 a n d OFL62,  in  low  d u r i n g summer  I would e x p e c t  range c e n t r e s l a t e r a l l y  some  This  elevations (Fig. 5).  of  and o t h e r s  them  If to  downwards  between summer and w i n t e r . Slopes were  not  VIII). and  on t h e s p r i n g and summer home r a n g e s as  steep as those  Exceptions  steep  sidehills,  gently  s l o p e d . No c l e a r were  winter  were t h e s p r i n g home r a n g e s  t h e summer home r a n g e s  ranges  on t h e i r  and  evident  o f OFL58  the winter trends  and  most  home r a n g e s  deer (Table  o f OFL58 and OFL60  OFL62  which  were  on  home r a n g e o f 0FL71 which was  i n the aspects of  (Table V I I I ) .  of  seasonal  A l l a s p e c t s were  home  represented  106  Table  V I I I . Slope  home r a n g e s .  and a s p e c t  near  t h e a r i t h m e t i c mean c e n t r e s o f  No c o n s i s t e n t t r e n d s a r e e v i d e n t .  Slope spring  0FL61 OFL68 OFL67  winter  spring  summer  winter  9  24  N-NW  SW-N  SW-W  8  22  S-SE  SW-SE  7  29  summer  ^1  3  Aspect  (degrees)  NW-W S-SE  OFL60  23  6  winter range  15  "  a u x i l i a r y winter range  24  10  0FL58  23  0FL62  4  * horizontal migrator  mm  -  "  0FL71 *  N-NE  W-S-E  NW-NE  E S-E  W-S-E  NW  10  NW  14  11  W-SE  W-SE  SW-SE  34  33  W  NW-W  NW-W  £1  108  in  s p r i n g and summer home.ranges. I n w i n t e r ,  located  on  aspects  varying  southerly  home  ranges  between  were  northwest and  southeast. Seasonal Figure the m  20.  movements o f r a d i o - t a g g e d In general, a l t i t u d i n a l  beginning in  of w i n t e r and occupy  elevation.  Their  spring  bottoms a d j a c e n t  to their  deer  high  moved  to  in  their  home  changed  substantially  between  the  deer  narrow  climate  a  seasons.  migrations  similar  to  In  Where  those  small  shifts  local  climate  distance,  attained  these  i n h a b i t e d low  such  and t h e wide main  and  650  i n valley  summer,  deer  horizontal valley  below  were u s u a l l y  Resident  in  moved downward a t  and made r e l a t i v e l y  tributary  made h o r i z o n t a l  local  ranges  between over  summarized  home r a n g e s  w i n t e r home r a n g e s .  t h e year  ranges  are  migrators  winter  elevations.  e l e v a t i o n s throughout  deer  valley,  differences  experienced  by  as  in  altitudinal  migrators.  3.5.  Indirect The  movements  corroborated population. individual the  deer  M e a s u r e s Of S e a s o n a l  by i n d i r e c t These  data  by  the  radio-tagged  deer  were  m e a s u r e s o f movements made by t h e d e e r  measures  movements  were  were  employed  b r o a d l y based  to  evaluate  i f  and r e p r e s e n t a t i v e o f  population.  Deer t r a c k s February  made  Movements  to July  were  observed  i n 1975 and from  along  October  were combined a n d t h e e l e v a t i o n  on  Mount  C a i n road  21.  The e l e v a t i o n  during  Mount  Cain  road  from  1975 t o May 1976. The  of the highest  deer  track  w i n t e r a n d s p r i n g i s shown i n F i g u r e  o f the h i g h e s t deer  t r a c k decreased  from  1200  109  Figure  2 0 . S e a s o n a l Movement p a t t e r n s o f d e e r  Arithmetic each  deer.  mean  c e n t r e s o f s e a s o n a l home r a n g e s  = spring, (s)  rinter. Cutover  i n the.study  = summer, (*T)  areas are stippled.  area.  are joined f o r  = winter,(A)  =  auxiliary  110  111  Figure. Cain  21.  road.  E l e v a t i o n of highest  deer  t r a c k observed  along  Mount  summer  spring  winter  1200H O  p  800  > U J  UJ 4 0 0 H  O #  O 1975 % 1975-1976  0 0  N  D  J  F  M  A  M  J  J ro  T I M E  ( M O N T H S ' )  113  m  in  early  elevation and  October  of deer t r a c k s  increasing  winter, and  The  700  V I I ) . In  elevation  of  spring,  with track  counts  night  increased  at middle  visits  t o about  of  deer  track  moved  elevations, moved.  appearances  50% i n t h e  tracks  altitudinal  transects the  transects  high  elevations by 0FL61  high  elevations  counts  indicate  showed  movements.  Since  (Section indicate after  deer  since v i s i b i l i t y ,  between t r a n s e c t s .  deer  km  -1  in  after  areas.  to  The  cannot  the s p r i n g  when  the  majority  a  first  in of  prevented  low deer  o f deer  make  vehicular  abundance o f d e e r The  within  magnitude  be compared between a l l  and t h u s t h e a r e a c e n s u s e d ,  However, w i t h i n  making  3.4.4).  majority  snow  relative  deer  wintering  the  the  deep  by  d u r i n g p e r i o d s w i t h s h a l l o w o r no snow. of  of  cutover  similar  when  when  night counts indicate  number  consistent  ( F i g . 21) and s i g h t i n g s o f d e e r  ( F i g . 21)  t h e y do n o t  that  increasing  ( F i g . 22) and were  counts  into  Night  travel,  to  showed  and h i g h e l e v a t i o n s o c c u r r e d soon  behaviour e x h i b i t e d  Although  winter.  throughout  with  22) i n t h e h i g h e r t r a n s e c t s may be c a u s e d  migratory  ( F i g . 20  in  i n t h e Mount C a i n s e c t o r  ( F i g . 2 1 ) . The f i r s t  observations  temporary  of  than those observed  progressed  snow c o v e r was r e d u c e d first  area  During  h i g h e s t o c c u r r e n c e s o f deer  s i g h t i n g s o f deer  spring  counts  (Fig.  the  2).  line  summer.  dates of f i r s t as  (Fig.  snow  h i g h e s t deer t r a c k i n c r e a s e d  Night counts conducted  elevation  with the descending  low e l e v a t i o n s i n t h e s t u d y  the  and on i n t o  i n November. T h i s d e c r e a s e i n  i n high elevations  were h i g h e r i n e l e v a t i o n  spring  the  m  coincided  snow d e p t h s  deer i n h a b i t e d  Table  tracks  to  transect,  the  differed  number  of  114  Figure  22.  Mean  number  deer  (vertical  bars)  Mount C a i n  s e c t o r . The number o f  above the  the  counted  of  95% c o n f i d e n c e  low e l e v a t i o n s i d e h i l l  deer  during  winter.  limits. forest  Transect  22-B was  t h a t was used  combined.  are  given  adjacent  intensively  to by  A was f u r t h e r f r o m t h e f o r e s t e d  and s i t u a t e d  herb s e r a i  limits  transects i n the  Transect  R i v e r . E l e v a t i o n s are i n d i c a t e d  E were t h r o u g h  road  counted  Davie  and  confidence  transects  range than  A and B were t h r o u g h  95%  night along  winter  Transects  B  at  and  i n the v a l l e y  conifer stages.  b o t t o m next  and i n c r e a s e f r o m serai  stages,  to  A t o E.  while  C,  D,  Data f o r 1975 and 1976 a r e  winter  spring  summer  winter  116  deer km  gives a relative  -1  trends  i n d e x of p o p u l a t i o n d e n s i t y .  between t r a n s e c t s show t h e  Seasonal  movement p a t t e r n s o f t h e  deer  population. The night  altitudinal  (Fig.  d e e r and  pattern  elevations, by  early  in  spring  deer  22)  deer  pattern of deer  was of  consistent  (see  densities  section  3.6).  early  tagged  ( F i g . 18)  year  3.6).  (Fig.  resident  and  date  of  as e l e v a t i o n  final  duration The  peak  with  increasing  moving  by  deer  use  migration  15 and  16) .  and  a result but,  o f deer  home r a n g e s  summer home r a n g e s . C o m p l e t e , o n e - s t e p  the  0FL61,  0FL67,  beginning  elevation  of  transects  OFL60, and winter, 22-D  OFL71  deer and  use 22-E  areas  t h a t had  their  10 and  decreased  short  (Fig.  to their  migration  made  of  represents  from  (Fig.  the  use i n  progressively  directly  (Figs.  deer  a result  ( F i g . 2 2 ) . T h i s peak  spring  radio-  progressively  22). F i r s t  a l s o was  of  radio-collared  a r e a was  rather,  lower  throughout  summer home r a n g e s  o f d e e r use  elevation  the  home r a n g e s . I n d i v i d u a l d e e r m6ved d i r e c t l y  by  use a t  of f o r e s t e d  elevations  i n an  areas  radio-tagged  deer  use  low  decreased  logged  of  by  ( F i g . 21  deer to t h e i r  their  low  (Figs.  probably not  i n intensity  from  in  deer  represented  increased  spring  visits  but  elevation  locations  increased  are  first  h i g h e l e v a t i o n s was their  and  at  ( F i g . 2 ) . At  i n spring  decrease  Some d e e r i n h a b i t e d  22)  areas  summer c o r r e s p o n d s t o m i g r a t i o n s  d e e r OFL62 and OFL58  The later  The  melt  o f low  the  logged  movements o f r a d i o - t a g g e d  were g r e a t e s t  a l s o e v i d e n t from  deer  in  snow a c c u m u l a t i o n and  e l e v a t i o n s by  (Section  with  summer. T h i s i n t e n s e use was  use  was  22). T h i s  later deer summer  spring  to  exhibited  12 t o in  20).  14).  the  At  high  decrease  117  coincided  with  elevations  the  first  decreased,  snowfalls.  deer  use  deer.  documented deer  This  downward  movements  deer  increased  ( t r a n s e c t s 22-A and 2 2 - C ) , i n d i c a t i n g some  As  a  movement  at  was  increase  greater during 3.6)  and  cutovers  in  numbers  winter than  elder  high  movement  consistent  by  with the  o f 0FL61, OFL68, and OFL67. The i n c r e a s e o f  differences were  at  low e l e v a t i o n s  downward  numbers a t low e l e v a t i o n s was s m a l l . The  small  use  was  that  i n the  between  deer  other  and  for  this  use o f f o r e s t s  seasons  transects  and had t a l l e r  reason  (see  section  in visibility  denser  was  (lower  vegetation  than  the h i g h e r c u t o v e r s ) . In  the  dispersion Shifts  22). from  of the  in  interval  in  (1)  deer  sector  was o b s e r v e d  t h e mouth  occurred  winter  use i n Croman  the v a l l e y .  over  beginning  Valley  was  mouth. As s p r i n g  o f deer  of  deer  on  the  population migrated  towards  Davie  River.  sector (Fig.  i n order of d i s t a n c e with  transect  23-A  of winter, deer  while  use  decreased  OFL60 and 0FL71 were c o n s i s t e n t  with  observed  These  shifts  through  the n i g h t  a l o n g Croman V a l l e y  f o r the winter horizontally  increased  further  the  Valley  i t  moved  spring  near  a l o n g Croman  sidehill  (2)  elevation  greatest during early  c o u n t s . 0FL61 d i d n o t move h o r i z o n t a l l y stayed  a narrower  p r o g r e s s e d , deer  Near t h e b e g i n n i n g  movements  ( F i g . 23).  range.  upper p o r t i o n s o f the v a l l e y  movements  occurred  i n t h e Mount C a i n  o f Croman V a l l e y  to a f o r e s t e d  The  also  i n F i g u r e 23 were a r r a n g e d  the v a l l e y  mouth.  ( F i g . 1 ) , s e a s o n a l changes i n  population  population  Transects  Deer  up  the  than  adjacent  near  Croman  in in  but  ( F i g . 1 0 ) . Most o f t h e Croman the  Valley, deer  south  population  118  Figure  23.  (vertical  Mean bars)  Croman S e c t o r . the  95%  number counted  of  deer  at  night  limits.  A t o D. E l e v a t i o n s  were  herb  serai  confidence  from  are: given  above  the_ v a l l e y  mouth  are indicated.  stages.  limits  along road t r a n s e c t s i n the  Distance  i n c r e a s e s from  combined.  95%  The number o f t r a n s e c t s c o u n t e d  confidence  through  and  Data  All  transects  f o r 1975 and 1976 a r e  winter  winter  spring  580 m  D  LL) LU Q  3  J  F  M  A  M  T I M E  J  J  A  S  ( M O N T H S )  O  N  290m A  D  120  corresponded t o patterns  o f snow m e l t i n s p r i n g and s n o w f a l l and  snowpack a c c u m u l a t i o n  i n winter.  3.6.  Habitats  Use  Seasonal  Use Of  O f U n l o g g e d And L o g g e d Day  and  classified  night  as  locations  being  o f age) d u r i n g  relative  u s e made o f t h e s e  percent  use  movement p a t t e r n forest  during  opposite use  cutovers  of  the  forested or cutover  three  habitats  was  occurred  during  show  two  types  some  while  cutovers  24 and T a b l e  were used  summer  their  use o f  each  The  season, use o f  deer  to  the  i n use o f f o r e s t e d and during  day  than  at  more a t n i g h t t h a n d u r i n g day use  of  d e e r was g r e a t e r  d e e r were c o n s i d e r e d  forested than  spring  and a l m o s t  decreased  together,  t h e day t h a n a t n i g h t ,  more f r e q u e n t l y a t n i g h t .  winter  seasonal  that  ( T a b l e IX) .  f o r e s t s was g r e a t e r d u r i n g  during  the  t h e day.  migratory  more  by m i g r a t o r y  When a l l r a d i o - t a g g e d  in  Within  I X ) . D a y t i m e and n i g h t t i m e  during  r e s i d e n t deer  of  similarities  night,  lowest  (up t o  f o r each  t h a n i t was d u r i n g  h a b i t a t s . F o r e s t s were used  were used  areas  day t h a n a t n i g h t , w h i l e  cutover  by  were  indicated  deer i n c r e a s e d  i n use of c u t o v e r s .  was g r e a t e r  deer  habitats  and  averaged  ( F i g . 24). Resident  Comparison o f t h e  (Fig.  seasons  deer  h a b i t a t s by d e e r t h r o u g h o u t t h e y e a r .  was g r e a t e r a t n i g h t  resident  radio^tagged  b o t h day a n d n i g h t between s p r i n g a n d w i n t e r .  trend  of forest  of  i n either  27 y e a r s  The  Habitats  D a y t i m e use o f  doubled  slightly  while  from  i n summer. that  use o f cutovers  forests  was  Use o f f o r e s t s  during  summer.  In  121  Figure each  24. P e r c e n t use o f f o r e s t e d season  (SP = s p r i n g ,  and c u t o v e r  SM = summer, WT = w i n t e r ) .  h i s t o g r a m s a r e d a y t i m e u s e , shaded Number o f d e e r f r o m cutover  habitats  which in  each  habitats  the  Unshaded  histograms are nighttime use.  percent  season  above t h e h i s t o g r a m s f o r f o r e s t e d  during  use  of  was c a l c u l a t e d  habitats.  forested  and  are indicated  P E R C E N T  123  T a b l e I X . C o m p a r i s o n o f use o f f o r e s t e d a n d c u t o v e r migratory in  forested  and  resident  deer during  h a b i t a t s more f r e q u e n t l y  habitats  summer. M i g r a t o r y than  were r e s i d e n t  deer  by were  deer.  Night time  Daytime n  migratory resident  181 58  forested  cutover  % in forest  n  forested  cutover  % in forest  162  19  89.5  81  26  55  32.1  kO  18  69.0  17  2  15  11.8  X= 2  12.6  ,  p  £.05  Fisher's Exact P r o b a b i l i t y Test p= . 0 5 9  125  cutovers,  daytime  use  summer, a n d i n c r e a s e d observed spring  in  through  comparable  Use  spring,  lowest i n  again i n winter, but not to the  intensity  spring.  was  At  greatest  night,  summer t o w i n t e r ,  use o f f o r e s t s i n c r e a s e d  Associations  Locations  of  radio-tagged  of use c a l c u l a t e d  serai  these  stage  habitats  nighttime  i s  not  selection covered  known,  insight  deer  where  were c o n s i d e r e d  and  were  preference.  with  cursory  appropriate. separately.  Since  cannot  type.  Regardless of h a i i t a t  do i n d i c a t e t h e v a r i o u s  estimates  a  association  Daytime  be  terms  stage  o f each  used d u r i n g  is Some  comparison  the l o c a t i o n s  of  o f the  calculated.  of the extent  and  C a u t i o n must be  the p r o p o r t i o n  through  preferences,  habitats  and  v a r i a t i o n i n the use of  i n t o p r e f e r e n c e c a n be o b t a i n e d  use  plant  association or s e r a i  preferences  deer  combined  frequency d i s t r i b u t i o n s i n  by e a c h p l a n t  habitat  Stages  f o r each f o r e s t e d  discussed  locations  study area  And S e r a i  (Fig. 25). I n d i v i d u a l  used i n i n t e r p r e t i n g these habitat  a  decrease.  Plant  and  from  w h i l e use o f c u t o v e r s showed  Of F o r e s t e d  frequency  during  of  habitat of  deer  d i f f e r e n t seasons.  Spring In  spring,  forested  plant  Western  Hemlock  a l l deer  associations Zone.  was t h e most f r e q u e n t l y The  Amabilis  associations  use d u r i n g  the  serai  stages  in  The V a c c i n i u m - S k u n k Cabbage used f o r e s t e d  Fir-Western were  or  b o t h day a n d n i g h t  Hemlock  next  most  habitat and  the  Coastal  association  during  Salal-Western  intensively  was i n  used  daytime. Hemlock forested  126  Figure  25.  stages  Percent  by  use  of f o r e s t e d  radio-tagged  histograms are  deer  d a y t i m e use,  plant  during  shaded  associations each  and  season.  l o c a t i o n s are  serai  Unshaded  nighttime  us^e-. ui  Number  of  deer  from  cutover habitats above  the  :  VS  Fern - Western  Red  = Western  ftT  =  for  =  SD  DW  =  habitats.  Skunk Deer  Fern  F i r - Twisted  L o g g e d , H = Herb, F = F e r n ,  AW  - Douglas-fir,  Immature  of  forested  are  Serai  S = Shrub, C =  SC  - Western  plant  =  = Salal  Stalk, Stages  MC :  Conifer.  Sword  Hemlock,  = Amabilis SW  and  indicated  Forested  Cabbage,  Plagiothecium,  = Salal  .  -  use  calculated  forested  Cedar,  Amabilis  Hemlock - C o p p e r b u s h  percent  was  Vaceinium  Hemlock -  W e s t e r n Hemlock, Hemlock,  the  i n each season  histograms  associations  WP  which  Fir -  =  Western Mountain  N =  Newly  127  summer  40' LU O tr LU CL  day  n= 7  night  n = 6  •  L  winter  40H  day  n >= 7  night  n =  Li  Q L  SC DWWP AW SD SW  C o a s t a l Western Hemlock  Zone  n —  7  20  VS  J  AT MC Mountain Hemlock Zone  .1 , rjll  N  H  Sera! Stages  128  habitats.  Deer  Plagiothecium for  Fern-Western  associations  or  Western  minor use,  Sword F e r n - W e s t e r n  o f any a s s o c i a t i o n s Use  night  of forested  than  similar the  had o n l y  and  each  Hemlockaccounting  l e s s t h a n 6% o f t h e d a y t i m e u s e . T h e r e was no d a y t i m e  Salal-Douglas-fir nor  Hemlock  during  i n that  most  plant the  associations day.  Mountain was  However,  associations  Hemlock  generally  the pattern  while  Fir-Western  secondary  Hemlock  use  and  was  Zone.  less  at  o f u s e was  t h e V a c c i n i u m - S k u n k Cabbage a s s o c i a t i o n  intensively  Amabilis  i n the subalpine  Red C e d a r  use o f  was used  made  of  Salal-Western  the  Hemlock  associations. In  cutovers,  conifer the  s e r a i stages,  shrub  than  s e r a i stage.  during  intensively the  the greatest  the  Use of s e r a i s t a g e s  day.  used h a b i t a t  The  herb  at night.  most commonly  stage  was t h e t h i r d .  logged  use  was  in  herb  and  w h i l e l e s s t h a n 5% o f t h e d e e r use was i n  second  serai  daytime  serai  was g r e a t e r  at  stage  the  most  stage  was  The c o n i f e r  was  serai  night  used c u t o v e r h a b i t a t ,  while the shrub  No use was  fern  s e r a i s t a g e s by r a d i o - t a g g e d  made  of  deer during  or  newly  spring.  Summer In  summer,  association plant  was a g a i n  associations  25).  This  was  mainly  resident was  daytime  still  high  use  greater in  the  use of the  than  the the  Coastal  Vaccinium-Skunk use  migrator  of  0FL71  Cabbage  forested (Fig.  association  ( F i g . 2 6 ) . However,  used t h i s a s s o c i a t i o n .  made o f A m a b i l i s  other  Cabbage  Western Hemlock Zone  Vaccinium-Skunk  due t o h o r i z o n t a l  d e e r OFL62 a l s o  of  In  summer,  F i r - W e s t e r n Hemlock, W e s t e r n  use  Hemlock-  129  Figure  26. P e r c e n t  stages during and  use o f f o r e s t e d  plant  summer by a l t i t u d i n a l  residents.  Unshaded  and  histograms  associations  and s e r a i  horizontal  migrators,  are  histograms a r e nighttime  u s e . Number  percent  and c u t o v e r h a b i t a t s  use o f f o r e s t e d  calculated  are  indicated  habitats.  Forested  plant  above  of  daytime  the  associations  deer  Hemlock,  AW = A m a b i l i s  WP = W e s t e r n  F i r - Western  Hemlock,  Hemlock - C o p p e r b u s h  N = Newly L o g g e d ,  i n each  the  season  for  was  forested  DW = D e e r  - Skunk Fern  -  Hemlock - P l a g i o t h e c i u m , SD = S a l a l  SW = S a l a l - W e s t e r n Hemlock, AT = A m a b i l i s MC = M o u n t a i n  which  VS = V a c c i n i u m  Cabbage, SC = Sword F e r n - W e s t e r n Bed C e d a r , Western  from  histograms :  u s e , shaded  .  H = Herb, F = F e r n ,  -  Douglas-fir,  F i r - Twisted  Immature S e r a i S = Shrub, C =  Stalk,  Stages :  Conifer.  130  altitudinal  day n = 4 night  n = 3  40-  20-  T  1  1  -,—q-  r  horizontal  604  day n * 1 night  rLU O tr LU  1  n = 1  40  20H  0-  -1—q— f* L  resident  60H  day n = 2 night  n = 2  40-^  20H  T  VS  r  1  SC DWWP AW SD SW  Coastal Western  Hemlock  Zone  AT MC Mountain  Hemlock  Zone  N  H  F  S e r a i Stages  S  4C  131  Plagiothecium, d a y t i m e use deer in  of  (Fig.  spring,  and  Salal-Western  these associations  26).  Two  were the  due  to a l t i t u d i n a l  similar was  to  that  associations Serai than  in  almost 5%  of  40%  Use  of  the  d a y t i m e use  elevations.  Thus, t h e  to  conifer  but  of  than  conifer  i n spring  a r e s u l t of  the  conifer  conifer  much l o w e r  the  i s not  lack  of  X),  w h o l l y due  had  suggesting  herb  spring.  of  the  f e r n s e r a i stage r e s u l t e d from  0FL71,  was  cutover c l a s s i f i e d  she  used  the  area  which  less  than  the  stages  not  have  resident and  high  conifer  Croman  of  summer.  in  serai  only  in  deer  spring are  stage.  than one  Creek  in  deer,  a f e r n s e r a i stage  near  the  decrease  l e s s i n summer  as  of  other f a c t o r s  the  s e r a i stage  summer  movement  spring  t h i s d e e r was  of  to  OFL62 i s a  of  by  same  in  migrators did  Daytime use  OFL71. A l t h o u g h t h e  i n use  in  old serai  f o r the  Use  reduction  the  t h i s d r o p i n use  responsible  used  and  s e r a i stage,  s e r a i stage during  (Table  was  associations  day  occurred,  altitudinal  s e r a i s t a g e by  i n summer  mainly  i n summer  plant  during  conifer s e r a i stage.  the  these  periods.  The  s e r a i stage  from t h e  with a c c e s s to Use  of was  t h i s s e r a i s t a g e i n summer. However, t h e  the  d e e r away  Zone  not  Mountain  Use  associations  i n summer. Some o f  at  of  and  25) .  forested  both  d a y t i m e use  was  use  of  25).  s e r a i stage  access  (Fig.  plant  night  conifer high  Stalk  s t a g e s were used i n d i f f e r e n t i n t e n s i t i e s (Fig.  resident  ( F i g . 26) .  were u s e d d u r i n g  spring  the  at  the  Most  used i n summer, b u t  M o u n t a i n Hemlock  of f o r e s t e d  less  from  Fir-Twisted  migrators  in spring.  generally  associations  subalpine  use  associations.  resulted  associations  i n the  Nighttime  plant  Amabilis  Hemlock-Copperbush associations  Hemlock  was more  132  Table X. Daytime use o f the c o n i f e r s e r a i stage by r e s i d e n t deer OFL62, during  s p r i n g and summer.  X =9.64, p<.05 2  Spring  Conifer Serai Stage Other Habitats  Number of Locations % i n Conifer Serai Stage  Summer  13  3  16  33  29  36 8  ,  134  intensively was  than  greater  the r e s t  i n shrub  of the cutover.  and h e r b c o v e r  T h i s stream  than  bank  were c e n t r a l  area  portions  of the cutover. Trends i n nighttime trends  in  greater  ( P i g . 2 5 ) . As i n d a y t i m e ,  serai  stage  s e r a i stage deer.  use, although  resulted  solely  was n o t used a t  This  spring. than  daytime  use o f s e r a i  Use o f h e r b a n d f e r n  s p r i n g , while  were  the i n t e n s i t y nighttime  from  in  with  serai  to  o f use was much  use  summer  the high  stages  use o f t h e s h r u b  similar  of  the  fern  ( F i g . 2 6 ) . The c o n i f e r  0FL71  night  c o n t r a s t s markedly  stages  by  radio-tagged  use i t r e c e i v e d i n  was h i g h e r  serai  stage  was  in  summer  lower.  Winter In w i n t e r ,  d a y t i m e u s e was g r e a t e s t i n A m a b i l i s  Hemlock  and  Salal-Western  Secondary  use was made o f  Plagiothecium, but  and  use o f t h e s e  two  bottom  the  Cabbage  winter.  Davie  Cain  of  the  Sword  River  Hemlock-  that of the  Fern-Western  association  was  type  used  during  subalpine  M o u n t a i n Hemlock Zone were  i n c o n t r a s t t o the high  former  Red C e d a r the  (Fig.  o f h a b i t a t f o r deer a r e a . The  valley 14). This i nthe  Vaccinium-  only o c c a s i o n a l l y during  T h i s low use c o n t r a s t s s h a r p l y and  into  n e a r Hoomak L a k e  received  winter  than  s e c t o r s o f the study  spring  2 5 ) .  F e r n - W e s t e r n Bed C e d a r a s s o c i a t i o n s ,  was n o t an i m p o r t a n t  Croman and Mount Skunk  Sword  Use  (Fig.  S a l a l - D o u g l a s - f i r , Western  was due t o 0FL71 when s h e m i g r a t e d  along  association  associations  h a b i t a t s was much l e s s  associations.  association  Hemlock  Fir-Western  with  summer.  The used  use t h e y  the  high  use i t  a s s o c i a t i o n s i n the incidentally  during  received during  summer.  135  Although  use o f f o r e s t e d h a b i t a t s a t n i g h t  than  that  in  daytime,  night  were s i m i l a r  than  use o f  o f herb  shrub  winter d i f f e r e d and  conifer  from  serai  that i n stages.  and f e r n  serai  stages i n winter  was  t h a t i n summer. The c o n i f e r  serai  s t a g e was  more  night  in  stages  r e c e i v e d about  not  less  t o t h o s e i n t h e day.  summer by t h e h i g h e r use  generally  t r e n d s i n use o f p l a n t a s s o c i a t i o n s a t  Daytime use of c u t o v e r s d u r i n g  Nighttime  was  winter than  exhibit  summer.  the  i n summer. H e r b ,  shrub,  used  and c o n i f e r  t h e same amount o f use i n w i n t e r  large  differences  apparent  less at  serai  and d i d  d u r i n g s p r i n g and  136  4.  Discussion  4.1.  Size  Of Home E a n g e s  Black-tailed British  Columbia  deer i n h a b i t i n g  coastal  and  migratory 1956),  winter  home  Home r a n g e s range  size  Harestad  was  with e x i s t i n g  deer t h a t  1963,  165  and B u n n e l l  Schoener  variability  ranges  (Dasmann  requirements  1968,  and s p a t i a l  of  North  home r a n g e  American  size  and  cervids  quality  100%  i s likely  in  a result  in  food  requirements  (McNab  p r e s s ) . The  of differences  abundance.  Sparrow  Home  (Cameron e t  and  1971, A l e x a n d e r 1973, The  decrease  is  ( i n press) t o t h e ruminant's (Nordon  et  a l . 1970)  o f food p r o c e s s i n g d u r i n g winter because forage  to  t h e y were i n summer. T h i s  e t a l . 1973).  by H a r e s t a d and B u n n e l l  of their  home  related  productivity  ( L e o p o l d e t aJL. 1951,  e t a l . 1973, P h i l l i p s  rate  behaviour  s i z e d u r i n g w i n t e r h a s been o b s e r v e d i n  Craighead  lower  deer  (1965).  i s  Bunnell,  Van B a l l e n b e r g h e and Peek  energy  size  and h a b i t a t  differences  1970,  apparent  Taber  f o r migratory  by K l e i n  Home r a n g e  Harestad  Springer  attributed  and  t o t h e e s t i m a t e p r e d i c t e d by  d u r i n g w i n t e r were s m a l l e r t h a n  decrease  1975),.  deer  i n summer and t h e mean  (in press).  home  i n a non-  concepts of migratory  ha , s i m i l a r  i n home r a n g e  body s i z e  lower  been o b s e r v e d  were p r o p o s e d  were l a r g e s t  the a n i m a l ' s energy  other  have  t h e e x i s t e n c e of s e a s o n a l home r a n g e s  are not c o n s i s t e n t  in  ranges  seasonal  summer, and w i n t e r , a l t h o u g h  population of black-tailed  for black-tailed  environments i n  were f o u n d t o have t h r e e p r i m a r y  ranges corresponding t o the s p r i n g , summer  forest  a l . . 1975,  and  o f reduced  Person  et  al..  137  The than  spring  both  the  productivity these  ranges  summer  and  relationships  small  located  home  spring  habitats  used  spring energy  would  mean a  nutrients  result  in  decrease  late  requirements,  sizes the by  home r a n g e s  of  food  during spring  the  deer  Harestad  over  t h e new than  the food during  amounts  in  from of  spring  but  used  : productivity  in  spring  energy  available the  summer.  t o summer and  other  home r a n g e  ( i n press) .  deer  and n u t r i e n t s ,  during  be  of growth, f a t  These changes  the  during the spring. i n this  to  d e e r ' s e n e r g y and  because  o f energy  o f home r a n g e s  ranges  and summer  on t h e s e a r e a s . F u r t h e r , n o t o n l y  availability  and B u n n e l l  the  growth  food  increased  time  and n u t r i e n t s  o f s e a s o n a l home r a n g e s energy  were  than  on s p r i n g  f o e t a l g r o w t h , and l a c t a t i o n .  and  home  quality  size  would be i n c r e a s i n g  i n the expansion  small  home range  in  than  explain  a s m a l l e r a r e a b e i n g needed t o o b t a i n  o f energy  summer  and  would be n e c e s s a r y d u r i n g w i n t e r .  to  requirements  deposition,  the  in  food  available  in  result  Energy  can also  The s p r i n g  t h e amounts o f f o o d a v a i l a b l e  would t h e d e n s i t y  nutrient  size  would be h i g h e r i n g u a l i t y  are similar,  decrease  lower  were s m a l l e r  ranges.  amounts  during winter, also  and n u t r i e n t s t h a n  home r a n g e s  would  greater  deer  i n w i n t e r . The abundance o f h i g h q u a l i t y  Although  The  home  ranges.  with  at t h e lower e l e v a t i o n s available  winter  w i t h home r a n g e  home  i n habitats  of black-tailed  summer  in  would from  The t r e n d s i n  study are c o n s i s t e n t model p r o p o s e d  with  f o r mammals  138  4.2.  Seasonal  Movements and N u t r i t i o n  Alterations respond  o f home r a n g e  t o environmental  maximum  limit  increased  to  changes.  the  extent  i s one way  altering  the  However t h e r e a p p e a r s which  position  to  home  of t h e i r  home r a n g e s . were  the b l a c k - t a i l e d  Black-tailed  deer  elevations  available. or  used  deer  u s i n g nearby  the  deer  other deer  migrated migrated  cutover  deer  cutovers  I n summer,  Although  nears  their  some d e e r  tributary  i s generally  used  home  selection  resulting  habitats  ranges  in  ranges  encompassed  the  home  ranges;  proximate  American  s e a s o n a l home r a n g e s  still  factors  ungulates, the  the  area, black-  during  deer  ranges.  these  of habitat  abundance o f f o o d  home  spring,  between  t o occur because  the r e l a t i v e by  more  accepted as seasonal f l u c t u a t i o n i n  are proposed  from  were  valleys.  and w i n t e r . Movements o f b l a c k - t a i l e d  seasonal  these  a t low e l e v a t i o n s b u t  ( O r r 1970, p. 2 4 ) . I n t h e s t u d y  different  area  bottoms  t o m i d - e l e v a t i o n s or high e l e v a t i o n s ; into  by  area.  they  home  o f use o f p o r t i o n s o f t h e i r  factor  deer  when  winter  remained  there are d i f f e r e n c e s  availability  study  moved down i n t o t h e v a l l e y  ultimate  in  is  altitudinal  the  habitats  t h e s e a s o n a l movements o f N o r t h  summer,  Another  h a b i t a t s on s i d e h i l l s a t  inducing  tailed  c a n be  changes  observed;  a  o f t h e s e a s o n a l movements  wintered i n forested  changed t h e i n t e n s i t y  food  size  In t h e study  population inhabiting  During spring  intensively.  other  t o be  h o r i z o n t a l m i g r a t i o n , and r e s i d e n c y . T o g e t h e r ,  patterns provide a general description  low  range  environmental  p a t t e r n s o f s e a s o n a l movement  migration,  of  by which d e e r c a n  ( C o v i c h 1976, H a r e s t a d a n d B u n n e l l , i n p r e s s ) .  way by w h i c h d e e r c a n r e s p o n d  three  size  supplies  Where  food  139  availability  fails  to  movements, o t h e r f a c t o r s the  relative  account  for  are suggested  favourability  of  patterns  which  could  of  seasonal  contribute  to  by b u r i a l  of  habitats.  Winter During food  w i n t e r , snow a f f e c t s  sources  burial  and  through  o f f o o d i t e m s by  downward subsequent snowpack  snowfalls. was  deep  availability. herbs  and  shrubs.  deer,  Similar  be  o f the f o r a g e and At high  enough  elevations by  late  Herbs  may  deer  yirginianus be imposed  a r e a . B e s i d e s h e r b s , the  shrub  the s h o r t e r  stems i s a l s o  available  (above  November  at  high  can  900 to  from  through  the  reduce  food  for  elevations  would  by  both  8 cm  of  white-  (Coblentz  1970).  deer  because  deep  the  no  observed i n enough  I f the e f f e c t o f  then  by  of  snowpack was  considered,  cause  m) ,  diets  t h e snowpack was  s h r u b stems.  The  burial  occurred  on b l a c k - t a i l e d  the  directly  digging  snow  deer.  permit t h e i r  (Zimmermann)  of  of  or  were c o v e r e d s u f f i c i e n t l y  evidence  bury  direct  t h i s f o r a g e component 0.  effects  study  r e s t r i c t i o n s on  This reduction i n a v a i l a b i l i t y  snow t o e l i m i n a t e tailed  locomotory  snow c a n  displacement  food a v a i l a b i l i t y  amount  to  bending of  be l e s s t h a n t h a t  food at  low  elevations. Food of  during winter a l s o  on  deer  mobility.  black-tailed  deer  i s hampered  of  snow  availability  the m o b i l i t y  deer  o f Bocky  Snow d e p t h s can  mountain  i n snow. I n t h e s e s p e c i e s ,  seriously  hampered  or  be  depends on t h e  at which  the  mobility  of  e s t i m a t e d from o b s e r v a t i o n s  mule  deer  and  white-tailed  snow d e p t h s o f o v e r 40  prevented  effect  locomotion  (Hosely  t o 60  cm  1956,  140  Loveless  1967,  shallower  snow d e p t h s  1959, of  Kelsall  Loveless  and  then  (Jones  greater  the  impeded than  increasing be  support deer  more  deer  migratory  would  occur  mean  for  black-tailed  which  their  mobility  depths  were  would be frequently  because  mobility.  the  Snow d e n s i t i e s  1975), so h i g h  This  in  increased  energy  that  reduced  i n a decreased  decrease  elevations  decrease  density  cost  of  snow-dependent winter  costs  deep snow  would a l s o  home r a n g e s of  of  effects  home r a n g e s  of for  1973). i n areas  available  obtaining food. For on  with  would be more f a v o u r a b l e t h a n with  to  o f snow t h a t would n o t  movement.  energy  area  degree  mobility  deer  summer  home r a n g e s ,  their  o f f o o d and t h e d e e r ' s  these  i n winter  their  impeded  (Jacobsen  deer,  Resident on  alees  f o r a g i n g i n a r e a s o f deep snow  increased  snow-free areas ranges  depths  Rices  their  result  an  efficiency  (Hepburn  h e i g h t s between 50 and 58 cm  (Fitzharris  and so r e s t r i c t  burial  and  seriously  t o have d e n s i t i e s  deep snow w o u l d r e s u l t  food,  (  Snow d e n s i t y d e t e r m i n e s  a f f e c t s deer  likely  mobility  The  at  while  t h a t snow  o f moose  i s a b o u t 37 cm. Snow  elevation  movements w h i l e  of  depth  mobility  relationship  chest  1970),  37 cm a t mid and h i g h e l e v a t i o n s o f t h e s t u d y  snow d e p t h  would  deer  same  average  most o f t h e w i n t e r .  which with  their  1975),  seriously  over  with  a^U  (1969) o b s e r v e d  white-tailed this  et  reduced  70% o f t h e c h e s t h e i g h t  movements, a s s u m i n g deer,  Gilbert  (25 t o 35 cm)  1967). K e l s a l l  g r e a t e r than  (Linnaeus))  1969,  foraging  shallow  the  snow o r  summer  home  cover. experience  during  lowered  food  densities  winter. This reduction  the c h a r a c t e r i s t i c s  o f food  and d i f f e r e n c e s i n snow d e p t h  and  p l a n t s on density  would summer between  141  forests  and  cutovers.  Use  of  cutovers  g r e a t e r i n summer t h a n i n w i n t e r the  ( F i g . 2 4 ) . I n summer,  f o o d i n t h e s e c u t o v e r s was i n t h e f o r m  herbs  were dormant o r dead  eliminated food  by r e s i d e n t  by even s h a l l o w  available  affected  by  g r e a t e r depths  food here  shallow  snow  than  nearby  observed  snow  densities  than  Winter  forested  depths  are  1975).  which  of  areas  have  which have g r e a t e r amounts  the  predominantly  of  their  cutovers  and  available  bottoms.  These  spring  to  S p r i n g home r a n g e s areas  i n w i n t e r , d i f f e r e n c e s i n snow home  deer  of  and  most  ranges their  deer  had snowpacks t h a t periods  snowpacks  in  winter  ranges  adjacent  sidehills.  S p r i n g home r a n g e s  Snow d e p t h s  would  Although not  also  i n the v a l l e y  t h e 37 cm d e p t h deer.  home  on  included bottom  f o r a g e would be c o v e r e d  the  cost of  located  than  home  low e l e v a t i o n portions  locomotion  ranges  to  did the  on  than  of  deeper  of black-  spring  home  g r e a t a s on summer home r a n g e s ,  on s p r i n g  in  accumulated  large  b u r i a l b y snow o f f o r a g e as  energy  were f r e q u e n t l y  estimated to r e s t r i c t  be  modify  were  f o rlonger  ranges  under  i n winter than  and p e r s i s t e d  tailed  energy  greater food  deer, being  g r e a t e r depths  than  less  costs f o r locomotion are  favourable  summer home r a n g e s  food  locomotion.  cutover.  or  are  At low e l e v a t i o n s  of resident more  shrubs  a r e a s and have h i g h e r  forested  between t h e w i n t e r and  valley  reduced  conditions.  with  amount  of  C u t o v e r s have snowpacks o f  most c u t o v e r s a n d e n e r g y  summer home r a n g e s  As  i s mainly  forested  conditions,  home r a n g e s  most  of h e r b s . In w i n t e r ,  availability  depths.  (Jones  habitats,  p o o r e r snow  their  was  snowdepths. F o r e s t e d a r e a s have more  since  costs f o r locomotion  lower.  and  deer  on  more  winter  142  home  ranges.  amount o f that and  On  food  available  available the  s p r i n g home r a n g e s , during  winter  h e r b component i n  would be  i n summer o r s p r i n g . G i v e n  h i g h e r c o s t s of locomotion  winter,  the  winter  home  ranges  on  much  the  less  than  absence of  herbs  s p r i n g home.ranges  would  be  the  the  most  during  favourable  habitats. although  d i f f e r e n c e s between i n d i v i d u a l d e e r  the l o c a t i o n s of s e a s o n a l characteristics. less  snowfall  during  present less  and  and  home r a n g e s were s i m i l a r  and  periodic  home r a n g e s .  winter . are the  snow c o v e r  more  food  habitats.  The  winter  available  and  the  Altitudinal  is  ranges are  migrations  t o cause m i g r a t i o n .  just  by  North  1975,  Capp  1968,  Hoskinson  and  Phillips  snow  (Edwards  Leege  food  other  is  seasonal is  low.  after,  winter  winter  the  5 cm  first  were  migration  of f o o d .  not was  This i s also  move t o  elevations  moved d i r e c t l y  to  their  line. winter and  e t a l . _ 1973,  Mech 1976,  is  d i d not  snow, b u t  of s n o w f a l l and  ungulates  the  of 3 and  availability  extent of  coincidence American  on  soon  Thus, t h e  o b s e r v a t i o n t h a t deer lower  of  amount o f  summer home r a n g e s t o  occurred  home r a n g e s w e l l below t h e  The  1963,  the  t o the  availability the  to than  o n l y h a b i t a t s where f o o d  o r more. S n o w f a l l s  response  below t h e  winter  cm  or  sufficient  in  12  from  of  implied  than  energy c o s t of l o c o m o t i o n  snowfalls  a direct  the  subjected  r a n g e s snow a c c u m u l a t i o n  available  home r a n g e s c o i n c i d e d w i t h ,  not  Abundance and  winter  snow  of s h o r t e r d u r a t i o n  a f u n c t i o n of both  snowpack. On  observed,  in their  W i n t e r home r a n g e s were i n a r e a s  were o t h e r s e a s o n a l food  were  and  m i g r a t i o n i s common Eitcey  LeEesche Hickey  1956, 1974, 1977).  Blood Sweeney Other  143  studies  observed  f a c t o r s such  1964),  Ashcraft  (1961)  occurring  up t o  occurrence  that  of  food  McCullough  several  weeks  (Cohen  migration reacting cover  to  either  snow  food  (Bergerud  not  an e a r l y  or  supply i s not  variable  o r premature  food  as w e l l as prevent  their  where  winter  may o n l y be which  migration  deer  response  their  supply  mean  inducing  highly  make  The  will  to winter  migrations  occur  s n o w f a l l s , d i s t a n c e s between summer and w i n t e r  to  environmental  food  t h a t a s n o w f a l l may o c c u r  s n o w f a l l deep enough  sufficient  migrations  ranges does  that  their  r a n g e s may be up t o 160 km. I n t h e s e of  1961).  ultimate factor  populations  1974). I n p l a c e s  the f i r s t  and S w i f t 1942,  snowfalls.  the  an i m m e d i a t e  covers  climatic  (Ashcraft  actual  before  the  with  observed  unpredictable  to the probability  their  before  i s  Ungulate  before  before  below t h a t on w i n t e r  include  1967).  (1964)  migrations  Adaptation  may  (Wright  availability  and  winter  correlated  and h u m i d i t y  water  availability  migration.  range  and  to densities  factors  migrations  as temperature  McCullough  reduced  winter  select  for  situations,  to  impede  deer  that  the p r o b a b i l i t y  migration are  may  be  sensitive  to  v a r i a b l e s which a r e p r e c u r s o r s o f snow.  In s i t u a t i o n s  where d i s t a n c e s  between  summer  and  winter  home r a n g e s a r e s h o r t , s n o w f a l l i s i m p l i c a t e d a s t h e t r i g g e r f o r migration. described  In  only  that occurs  snowfalls  (Dealy  in  deer  Although  i n late  area  i s  s e v e r a l months  1959,  anomaly, n i g h t c o u n t s Changes  one  McCullough  were  abundance summer t h e r e  made were  a  winter  m i g r a t i o n o f deer  before  the  1964).  In  only  on  the  interpreted  first this summer  as  was a d e c l i n e i n d e e r  winter apparent range.  migrations. numbers,  a  144  migration  to  the w i n t e r range  (1964) was u n a b l e  t o d e t e c t i n c r e a s e s i n deer  p o r t i o n s of h i s study population all  a r e a , and i n t e r p r e t e d  a s an e a r l y  other studies i n  winter,  snow  migration  Rempel  i s  et  areas  where  1932,  a l . 1952,  as  selection  f a c t o r s which i n d u c e  there  the  Dixon  abundance i n o t h e r  the decline i n  the  would  is  snow  initiator  1934,  Gruell  1977). D e p e n d i n g on  movements,  McCullough  m i g r a t i o n t o the winter range.  reported  (Eussell  Longhurst  was n o t d e m o n s t r a t e d .  1958, risks  Moir  and  f a v o u r deer  the  et  benefits  of  are necessary  trapped  on  loss i n fitness long  imposed  as i s p o s s i b l e  may be s u f f i c i e n t  by n o t s t a y i n g  (Cohen  these  were s e n s i t i v e t o  p e r s p e c t i v e . Over l o n g d i s t a n c e s , t h e c h a n c e s o f summer range  winter  a l . . 1951,  foraging  the  the  1976, B e r t r a m and  that  m i g r a t i o n s before they  In n e a r l y  during  of  Leopold  deer  from  a  being  t o outweigh t h e  on t h e summer  ranges  as  1967).  Spring During free,  and  locomotory would  spring, thus  when w i n t e r and s p r i n g  not  restrictions  uncovers  Snow m e l t s new  sources  w i n t e r home During  to by  the snow,  of a v a i l a b l e  on s p r i n g home r a n g e s  with higher d e n s i t i e s  burial  home  food than  o f f o o d . Use o f t h e s e of food  o f f o r a g e and  spring  early  a r e snow-  in  spring  ranges  winter  home  spring  and  home  ranges  r e s u l t s i n movement o f  deer  from  home r a n g e s  be c o v e r e d  ranges.  spring,  w i t h snow. L o c o m o t i o n the  o f deer  have g r e a t e r d e n s i t i e s  ranges.  their  subjected  home r a n g e s  summer  would s t i l l  would t h e r e f o r e be impeded,  snowpack c o u l d be l e s s  than  and  although  the snow-free h e i g h t s o f s h r u b s .  145  the  snowpack  on  summer  stem d i s p l a c e m e n t . shrubs  result  ranqes home  of  this  their  in  densities that  of  shrubs  because o f  consequent  a v a i l a b l e food  would be l e s s  than  burial  of  on summer home  those  on  study, l o c a t i o n s  and  Taber  as s p a t i a l l y  describe  spring  drifts"  spring.  Some a u t h o r s s t a t e  certain  magnitude  assumption  has  from of  w i n t e r home  results  were  different  t h e deer  or imply  that  to  of spring  because  et a l .  1952).  This  observed. studies  o f the deer  of i n d i v i d u a l  when Lack  probably  s e a s o n a l movements have been d e t e r m i n e d  locations  do  m i g r a t i o n s must be o f a  by e a r l i e r  changes i n t h e s e a s o n a l d i s p e r s i o n than from  but  discussions of "seasonal d r i f t s " ,  home r a n g e s  home  population during the  ( E i n a r s e n 1946, L o n g h u r s t led  results  ranges,  s h o r t d i s t a n c e s e a s o n a l movements were  recognition  rather  i n spring  (1956) who do n o t r e c o g n i z e s p r i n g  different  "seasonal  actually  o f deer  l o c a t i o n s i n winter. T h i s c o n t r a s t s with the  Dasmann  from  covers  ranges.  ranges  of  still  T h i s d i s p l a c e m e n t and  during spring  In from  ranges  usually  population,  animals.  Summer Energy (McNab  i s a major f a c t o r  1963, S c h o e n e r  t h e use o f h a b i t a t s predict of  that  available  Schoener need are  1965, Jarman  energy  linked  a primary (Blaxter  1974).  o f home  Foraging  should forage i n h a b i t a t s is  greatest  1971, Pyke e t a l . 1977).  i s  the s i z e  range  1968, H a r e s t a d a n d B u n n e l l , i n p r e s s ) and  (Klein  animals  influencing  Oldemeyer  a n d P i a n k a 1966,  ruminants,  one t o which r e q u i r e m e n t s 1962,  where t h e d e n s i t y  (MacArthur  For  1974)  models  the  of other and  the  caloric  nutrients available  146  energy 1957,  in  forage  Hardison  i s  indicative  1959, E e i d  1968)^  of i t s n u t r i t i v e  density in  movements  should  of available  habitats  used  seasonal habitats o f energy  place  energy in  ranges, deer  previous  s h o u l d be b a s e d  of  However,  digestible  Rochelle  ( i n prep.)  Digestible  dry  energy  with r  1971).  This  ruminants  2  digestible  dry  on t h e  energy  matter  and used  matter  i s  than  availability  were not a v a i l a b l e  to  were  of  proportional  supported  1973,  Hebert  evaluate  are usually  low  1969) . B e c a u s e o f t h e s e  home  ranges,  summer, t h e amount  t o d e e r on t h e i r  digestible  and  of energy  s e a s o n a l home r a n g e s  of  digestible  dry  dry matter per l o c a t i o n home  et a l . of  wild  Digestible  dry  ranges  in  crude  i s  of  each  fibre  interrelationships  dry matter of the a v a i l a b l e  o f energy and n u t r i e n t s a v a i l a b l e  density  energy.  to d i g e s t i b l e  studies  1973).  from  p r o p o r t i o n a l t o the p r o t e i n content o f a f o r a g e ,  of digestible  During  digestible  by  i s  area.  available  correlation  and H a r r i s  the density  place  directly  f o r the  deer i n t h e study  (Rittenhouse  (Ammann e t a l .  (Crampton  seasonal  relative  values  in  since forages high i n protein  the  would be g r e a t e r  of .98 (Moir 1961) and .93  i s also  density  h a b i t a t s where t h e  season. Thus s e l e c t i o n o f  major f o r a g e t y p e s used by b l a c k - t a i l e d  used  that  and n u t r i e n t s .  Estimates  matter  i t follows  in  and n u t r i e n t s  the  (Swift  G i v e n t h e r e l a t i o n s h i p s between  e n e r g y , f o r a g i n g b e h a v i o u r , a n d home seasonal  value  food  was  an e x p r e s s i o n o f t h e  t o deer. and n u t r i e n t s  available  was e s t i m a t e d i n t e r m s  matter.  The  mean d e n s i t y o f  was c a l c u l a t e d deer,  of  for the  three  under c o n d i t i o n s o f t h e  summer f o o d s u p p l y f o r t h e a p p r o p r i a t e h a b i t a t  types  listed  in  147  Table  I I . Mean  and  night  day  and n i g h t  a  mean  locations  calculated  in  density  and O c t o b e r  digestible  significantly  dry matter  (Table  d e e r was t h e  used  had i t  This  (List  length of  1966),  and  dry matter per l o c a t i o n  matter  < .05)  winter  in  ranges  use  Densities  of  during  summer  habitats  on  the  situated density of  contained  less  this  i t s winter  abundant  deer  would  home  in  Western  in have  range  during  and t h e l o w e r  density  d r y m a t t e r f o r h i s summer home r a n g e  during  home  were  one o u t o f t h e s e v e n r a d i o - t a g g e d  than  stayed  (p  d e e r , OFL62, was a r e s i d e n t  greater  association  their  habitats  encountered  of d i g e s t i b l e  than  For only  dry  summer.  greater  XI) .  digestible  spring  of  a c c o r d i n g t o average  l o c a t i o n s o f r a d i o - t a g g e d d e e r i n summer  digestible  his  between June  d r y m a t t e r f o r t h e day  ( T a b l e XI) .  habitats  summer  were w e i g h t e d  a t 50°N  daily  The  densities of digestible  Hemlock  -  was  due  to  Plagiothecium  summer t h a n i n w i n t e r .  o f d i g e s t i b l e d r y matter a v a i l a b l e ranges  and  different  (p < . 2 0 ) .  Three o f t h e f o u r m i g r a t o r y deer encountered  greater  densities  of  summer home  ranges  have e n c o u n t e r e d had t h e y s t a y e d on t h e i r  spring  digestible  t h a n they  were n o t s i g n i f i c a n t l y  on summer  dry matter  would  by moving t o t h e i r  home r a n g e s f o r summer. The f o u r t h resident of  deer  digestible  one  resident  density have  used  of digestible  been  h a b i t a t s i n summer t h a t  dry matter deer,  migratory deer  had l o w e r  t h a n h a b i t a t s t h e y used  t h e sample s i z e  o t h e r two d e e r i n c l u d e d  in  of l o c a t i o n s  d r y m a t t e r on t h e s p r i n g  overestimated. Part  and  o f the s p r i n g  the Vaccinium-Skunk  Cabbage  two  densities  spring.  In  was s m a l l and  home home  the  range  may  ranges o f t h e association.  148  Table  X I . Mean d e n s i t y o f d i g e s t i b l e  available ranks  d u r i n g summer on s e a s o n a l  dry  matter  home r a n g e s .  per  Wilcoxon  test. H :  summer < w i n t e r  H„:  summer < s p r i n g ; T=6, n=6, p<.20  H :  spring < winter  0  0  location  ; T=1, n=7, p<.01  ; T=3, n=6, p<.05  signed  149  Food A v a i l a b l e On Seasonal Home Ranges During Summer ( k g d i g e s t i b l e dry matter ha ) Winter Spring Summer Range Range Range  144.9  106.1  208.5  166.2  503.6  179.8  99.2  256.9  142.8  0FL71*  163.8  326.2  124.6  OFL58  288.8  210.0  172.0  0FL62  190.6  126.4  142.8  0FL61  360.4  OFL68 OS  o EH  OFL67  197.8  « (—I  En Z  0FL60  w Q M  CO  w PC  * h o r i z o n t a l migrator  150  This is  association  small  has a high  i n area  associations  .  spring elk  deer  o f d i g e s t i b l e dry matter but  relative  to  Migration  o f d e e r away from t h e V a c c i n i u m - S k u n k  Cabbage a s s o c i a t i o n resident  density other  and r e d u c t i o n  may  result  in  from  a n d use o f t h i s h a b i t a t  types  use  the  of  food  feeding  by  available i n spring  deer  the year  density,  deer  leave  predation. the  they  i n spring  habitats  density  and f o r a g i n g  i n June adding f u r t h e r t o the prey  time  (Orsus americanus P a l l a s ) because  of  predators to  on t h e s p r i n g  their  to spring  home  than  ranges  personal are tied  communication) to  low  dependence on v e g e t a t i v e r a n g e s would i n c r e a s e  summer home r a n g e s . I f  and i n t o t h e much l a r g e r  was s n o w f r e e , t h e r i s k  lower  prey d e n s i t i e s  of predation  (Tinbergen  why  habitats  et a l .  during  r a n g e s . Not  to  predators.  summer  ranges  t o den s i t e s and b l a c k  elevation food.  the r i s k  deer  on  b u t fawns a r e  elevation  d e e r i n h a b i t i n g t h e s e a r e a s and make s p r i n g  favourable  explain  range  available  would be l e s s on h i g h  Scott,  reduce  e f f i c i e n c y , then  ( Canis lupus Linnaeus) are t i e d  (B.  could  i n deer s p r i n g  born  this  habitats  t h e i r hunting  a r e t h e deer c o n c e n t r a t e d  because wolves  and  and go t o summer home r a n g e s , i s  would f o c u s t h e i r h u n t i n g  of predation  concentration  habitats.  only  Risk  Roosevelt  and e l k may r e d u c e t h e amounts o f  I f deer p r e d a t o r s s e l e c t e d o f prey  by  of deer i n  by  potential factor, that could  the spring  basis  spring  another  plant  habitat  concentration  throughout  While the e f f e c t of f e e d i n g food  forested  this  { Cervus elaphus r o o s e v e l t i Merriam). T h i s  associated  it  of  at  bears  habitats  Attraction of of  predation  home r a n g e s  moved  off  less  spring  a r e a o f summer r a n g e when may be r e d u c e d 1967).  because o f  151  Movement f r o m the  melting  summer  of  the  ranges.  immediate  but  spring  For  t o summer home  snowpack migratory  o f ground  ranges  prior  appear.  to  and  white-tailed  are proposed in  Eempel  Jli. c a l i f o r n i c us  for  the  1974,  Davies  reconnoitre  are  nutrient 1930,  These  their  summer  their  studies content  al.  1932,  1952).  important  was  1973).  although  reported  visits  and  mule  and  of  changes  ( S m i t h and  Sweatman  black-tailed  deer  p r o v i d e a mechanism  can be o c c u p i e d when  they  p r o v i d e d e e r w i t h an  up-  their  summer home  ranges.  as f a c t o r s  i n m i g r a t i o n s o f mule d e e r  1934,  (1976) habitats  both  Dixon  t h e abundance o f f o o d and i t s  Leopold et a l ^  nutritional  factors  1951,  are  i t also  are  1972,  as t h e i n d u c i n g a g e n t  protein  is  necessary  facilitates  (McLean  Longhurst  suggested  ( D e a l y 1959,  for  increased  growth  forage  B a i l e and F o r b e s  of energy  1974).  (Jones  Relative  as the  Hebert  and  body  digestibility  thus the b e n e f i t s of high p r o t e i n  mainly i n terms  by deer  Mech  implicated  Where  f o o d i n t a k e and  Ullrey  ranges  home  were o b s e r v e d i n  a n i m a l aware  suggest that  summer home r a n g e s  proposed  maintenance,  ruminants  summer  i n c a u s i n g s e a s o n a l movements, p r o t e i n c o n t e n t i n  forage  and  bare  to suboptimal  habitats  o f c o n d i t i o n s on  the f i r s t  been  by H o s k i n s o n  home  not  California  t o keep an  visits  with  was  t o the  have  for  of p o t e n t i a l  1977).  Russell  visits  and  and  more f a v o u r a b l e . S h o r t t e r m  Early  movement  weeks a f t e r  deer. Occasional v i s i t s  to-date assessment  et  (Canton))  supply  through which  deer  (1977)  as f u n c t i o n i n g  food  this  permanent s e a s o n a l o c c u p a n c y  some m i g r a t o r y b l a c k - t a i l e d  (Pli  deer,  S h o r t term  coincide  t h e u n c o v e r i n g o f f o r a g e on  o c c u r r e d up t o s e v e r a l  patches  Bertram  and  ranges  diets  to  Verme  and  availability  of  1972,  152  energy  and  nutrients  animals  depend  on b o t h  Studies r e l a t i n g (Dealy  on  1959,  the  seasonal  Klein  seasonal  home  quality  and  1965,  Hebert  1973) but  was  the  assessed,  behaviour  cannot  cannot  singled  be  quantity  and  digestible matter  to  In  out  matter  do  of  interior Island,  where  the  alpine  principal  areas.  These  c l a s s e s of forage  longer  p o r t i o n of the  1972). their  Both  would be If forage  in  as  deer  would  be  of the  study are  their  nutritive  deer  do  of herbs  between a l t i t u d e s  dry  in  nutritive in  drier  Vancouver  and  shrubs.  quality  over  grasses and  less on  plants  a  (Cook  shrubs  and  nutritional  the coast  than  areas.  for  the  area,  then  the  forage  herbs  forage  from  Both  moisture  i n the  deer  the  expected  and  a r e a on  would r e s u l t  i n the study  black-tailed  wet  quality  responsible  forage  digestible  much as i t d o e s  the  only d i f f e r e n c e s i n q u a l i t y ,  tailed  are  succulence  in interior  the  respond.  high  observed  were  d e n s i t y of  p h e n o l o g i c a l c y c l e than  t h e summer-long  prolonged  difference  maintain  migratory  i n migration.  differences  In  abundance  combine t o g i v e a d e n s i t y o f  senescence  forages of  of  forage  nutritive  food  value  summers  differ  forage.  advantages of  appear t o  altitudinal  f o r a g e would n o t and  deer  of  content of  since  variable  i t i s this  cause e a r l y  elevations,  content  and  migratory  have f o u n d  nutritive  of t h e f o r a g e  areas not  of  as the d r i v i n g  which b l a c k - t a i l e d coastal  low  made. The  quality  dry  deficiencies at  be  evaluation  of  quantity  movements t o p r o t e i n  d i s p a r i t i e s between s e a s o n a l r a n g e s , not  ranges  and  quantity,  of  s e a s o n a l movements o f several  observed  were e x p l o i t i n g  not  major  p a t t e r n s of  the  black-  differences movements. I f  a phenological state  of  their  153  forage as  i n order  t o maximize t h e i r  p r o t e i n , then  a slow  Deer would be e x p e c t e d move the  upwards best  in  i n t a k e o f some n u t r i e n t , s u c h  progressive  migration  to leave their  elevation or into  would  spring  home  tributary  was  deer  their  from  process  over  not observed.  The f i n a l  ranges  This  and  migration  pattern  of  of b l a c k - t a i l e d  s p r i n g t o summer home r a n g e s was n o t  many d a y s , b u t o c c u r r e d  observed.  valleys following  p h e n o l o g i c a l c o n d i t i o n of the forage.  movement  be  i n a period  a  gradual  of less  than  a  day. If black-tailed some  phenological  population stage be the not  deer  stage  changing observed.  order  forage,  within  the  some  and r e m a i n  home r a n g e s were a t  would  spring  migratory  during  remained  the high  If  phenological  stage  a d v a n t a g e s would  of  exist  on  whose  their  low  e l e v a t i o n h a b i t a t s were observed  system they  t h e h a b i t a t made a v a i l a b l e  o f t h e snowpack.  i n order  their  was  observations  t h e s p r i n g r a n g e s where  o u t over  the melting  d e e r were m i g r a t i n g  left  to  low e l e v a t i o n s i n t o m i d -  elevations,  and s p r e a d  summer w i t h  This pattern  t h r o u g h o u t summer. O t h e r d e e r ,  deer  this  i n response t o  there  behaviour,  deer  be e x p e c t e d  move  ranges u n t i l  were c o n c e n t r a t e d  the  deer  high  from  exploit  where  and n i g h t c o u n t  s n o w f r e e and p l a n t growth h a d begun. Under t h e of  area  t h a t moves a l t i t u d i n a l l y  Both r a d i o - t e l e m e t r y  to  then  p h e n o l o g i c a l s t a t e of the forage.  that  elevation  their  in  On Mount C a i n , t h e p o p u l a t i o n  in a horizontal strip  elevations  of  would be c o n c e n t r a t e d  occurs.  indicate  were m i g r a t i n g  to  forage,  f o r migratory  exploit then  deer,  would be s e l e c t e d a g a i n s t . Hence, a t h i r d  a  particular  since  nutritional  non-migratory  expectation  deer  i s that i f  154  black-tailed factor,  deer  were  responding  the population should  resident  deer  would  occur.  be  to  primarily  than  explaining others, of  nutritional  some  security  factors. home  animal's  fitness  that  factors  focuses  By a d d r e s s i n g  and  Seasonal  being  by  parameters  permits  optimal  external  social  context out  vital  exploitation,  invoked f o r  are  environmental  structures within  arise  of  which  functions  including  avoidance,  o f home r a n g e s r a t h e r  and G e i s t  predictions  than of  argue provide  primarily:  distribution  (1974), deer.  territories  Geist  ungulates  they  species  in  that an them  resource  m a t i n g , and r e a r i n g o f young.  movements o f b l a c k - t a i l e d  of  and  the i n d i v i d u a l s comprising  organization of ungulates (1974),  seasonal  i n f l u e n c e d and  variables  because  the  organization  systems  e t a l , . 1976). C r o o k e t a l . (1 976)  predator  social  explanation of  complex  to  seasonal  assessment  Organization  importance o f the s p a t i o - t e m p o r a l  Jarman  model  deer.  The  (1974),  for  n u t r i e n t s , and  factors,  a more g e n e r a l  societies  (Crook  particular  on e n e r g y ,  by  deer.  invoked  availability  these  Movements And S o c i a l  Mammalian modified  few  ranges i n terms o f the b e n e f i t s and c o s t s t o the  movements made by b l a c k - t a i l e d  confirm  migratory  numbers o f r e s i d e n t  m i g r a t i o n s and f o r a g e  movements  seasonal  carry  nutritional  my a p p r o a c h c a n be used a s t h e b a s i s f o r a s i n g l e  seasonal  4.3.  a  This expectation i s not confirmed  my o b s e r v a t i o n s . T h e r e a r e s u b s t a n t i a l Rather  solely  (1974)  of  resources  i s d i s c u s s e d by E s t e s  and i s r e l e v a n t t o t h e In t h i s by  study,  black-tailed  concerning  environments  t h e use  with  deer  the s p a t i a l seasonal  155  fluctuations i n availability  of  resources.  Group s i z e i s one o f t h e p r i n c i p l e s o c i a l used  to  describe  "chaparral" most  habitats,  o f t h e year  mainly the  organizations  b l a c k - t a i l e d deer  (Dasmann and T a b e r  o b s e r v e d . The " c h a p a r r a l "  closed  habitat  correlation Forested  by  Jarman  conifer  so s m a l l  deer  (1974) ,  stage  mainly  in  have  little  serai  stages),  1974,  Hirth  by  lateral  larger  than  further  seasonal  relationships  of  males.  During  (1974).  groups as  a  a  negative  the  habitat.  are  also  In  g r o u p s o r were  closed  the  study  solitary,  (1956) and a g r e e  In  cutovers  which  and  use o f t h e more  spring  group  dense h a b i t a t s  and summer  open should  sizes (Jarman  habitats result in  seasons than i n winter.  aspects  of s o c i a l  These  behaviour  such  and a c t i v i t y .  movements o f b l a c k - ' t a i l e d  influences  out  habitats  the visually  a f f e c t other  where some members spacing  in  s i z e d aggregations i n these  The  consist  observed  would be more open  deer during  interaction rates  groups  classed  of  for  (newly l o g g e d , h e r b , f e r n , and s h r u b  1977). The i n c r e a s e d  a g g r e g a t i o n s would as  who  be  by Dasmann and T a b e r  cover  the h a b i t a t  black-tailed  larger  would  small  w i t h t h e p r e d i c t i o n s made by Jarman  groups  and more m a l e - f e m a l e  habitat  serai  s i m i l a r t o those described  be  adult  g r o u p s s i z e s would be e x p e c t e d .  occurred  would  small  between g r o u p s i z e v i s u a l d e n s i t y  and  habitats area  form  1956). T h e s e  more m a l e s become s o l i t a r y  variables  (Crook e t a l . 1 9 7 6 ) . I n  o f m o t h e r s and young o r up t o t h r e e  rut  are  social  systems  on t h e i r  social  deer  organization.  are migratory, the seasonal deer  because  of  would the  affect addition  would In a  population  concentration  stable and  impose  and  hierarchical  loss  of  social  156  partners.  Seasonal  summer home r a n g e s would  result  established the  new  Even  d e e r some s o c i a l  on low  remaining  social  The  rut  1970) , and shuffle  and  so  occurs  the  deer  (Kucera  1978).  result  i n a less  transient  the  stable  deer  habitat  use  conceptual  Taber  early  winter  the  loss  of  between  the  December  migration  resulting  in  the  study  The  area  relationships  from  as i n r e s i d e n t Miller  the  among  would  The  winter  the  system  social  rather  (Thomas  period.  non-migratory  male h i e r a r c h y .  (1956) and  more  and  M i x i n g o f m i g r a t o r y and  by  A  resident  the  relationships  r e p e r c u s s i o n s on t h e mating  such  Model Of  in  t h e dominance h i e r a r c h y  social relationships  4.4.  would r e s u l t  population  hierarchical  Dasmann and  by  be  summer  would o c c u r . M i g r a t i o n away from  November  with  direct  of b l a c k - t a i l e d  to  the  areas i n h a b i t e d  would a l t e r  in  would a f f e c t have  After  have  on  were  partners.  overlaps  of  migration  this  spring.  would  deer encountered  e l e v a t i o n s by m i g r a t o r y d e e r  some p a r t i c i p a n t s and  valleys  relationships that  relationships  elevation  disruption  tributary  d u r i n g w i n t e r and  with the u n f a m i l i a r  range.  of the p o p u l a t i o n to  to  i n the breaking of s o c i a l  migration,  established  low  a t high e l e v a t i o n s or  between i n d i v i d u a l s  summer  home  movements by a p o r t i o n  males of  deer  deer would  organization be  based  than  on  populations  on  stable  described  (1970).  S e a s o n a l Movements succinct by  statement  black-tailed  model  o b s e r v a t i o n s of deer  (Fig.  27).  movements  o f t h e s e a s o n a l movements  deer This and  can  be  model the  made is  factors  through derived proposed  and a from to  157  F i g u r e 27.  Model of seasonal movements and  habitat s e l e c t i o n .  f—  Benefits To Fitness N A T U R A L S E L E C T I O N FOR  - ENERGY quantity and quality of food,  GREATER  FITNESS  requirements, expenditures.  HABITATS  HABITATS  AVAILABLE  SELECTED  HABITATS USED  Habitat Evaluation - experience - habitat predictability -  Seasonal  exploration  - natural selection for surrogates of fitness  1  Movements  •CD  Costs To Fitness - PREDATION security cover, mobility, other prey.  Seasonal Environmental Changes depth and density of snow, growth and dormancy of vegetation.  159  elicit  these  movements.  habitat selection this in  The  model i l l u s t r a t e s  by b l a c k - t a i l e d  deer  in  the  model, s e a s o n a l movements a r e a r e s p o n s e the  relative  number o f h a b i t a t s  favourability  the process of  study  area.  In  by d e e r t o c h a n g e s  of different  habitats.  Given  a  which a r e a v a i l a b l e t o d e e r , t h e use o f t h e s e  h a b i t a t s c a n be e v a l u a t e d on t h e b a s i s o f t h e b e n e f i t s and c o s t s to the user's f i t n e s s . represented  by a s i n g l e  parameter  integrates  weather,  competition,  within  The b e n e f i t s t o f i t n e s s  a particular  parameter,  the  effects  habitat.  security  the  impacts  c o v e r , and p r e d a t o r  Among h e t e r o g e n e o u s some w i l l  based  behaviour, deer  changes  (Bouckhout  long  tailed  may  also  patterns  behaviour have been  costs occurring  in  express  the  a h a b i t a t and prey  density,  avoidance. available  to deer,  others. This f a v o u r a b i l i t y  even d i r e c t  The e v a l u a t i o n for  evaluation.  the  is  c a n be  appropriate  Experience could  imminent  environmental  No c o n s c i o u s a n t i c i p a t i o n i s n e e d e d a s  appropriately contribute  deer, t r a d i t i o n  resident  alterations  w i t h a means t o a n t i c i p a t e  as deer respond  Tradition  of  habitats potentially  1972).  requirements,  of predation within  experience, s e l e c t i o n  or p o s s i b l y  provide  This  fitness  i n terms of the deer's f i t n e s s .  on t h e d e e r ' s  acquisition.  nutrient  The c o s t s t o  be more f a v o u r a b l e t h a n  evaluated  of  are  l o c o m o t i o n , and f o r a g i n g  e f f e c t s o f changes t o t h e r i s k incorporates  n e t energy  of a h a b i t a t  to the environmental  to this  "experience". In b l a c k -  probably helps t o maintain  i n individuals established.  signals.  once t h e i r  migratory  or  s e a s o n a l movement  160  Habitat  evaluation  explorations occurring visits  t o summer and of these  those  Although  they  fitness,  and  made  and  could  w i n t e r home r a n g e s  are  the  most  explain prior  t h u s more t h a n one  patterns of habitat  would  type  be  benefits  and  be o b s e r v e d  are e v a l u a t e d , deer  select  and  habitats  would  o f snow and  In  study  area these  favourability  as a n o t h e r  another  as  accrued through  is  explanation  seasonal  different  migrations habitats  by  i n the  of  the  would  habitats.  selection  would  the  food be  more  As  the  use  and  cover  thus  the  a  sources. change  habitats  o c c u r and  of  primarily  food  p r o c e s s e s a r e s e a s o n a l and  a  deer  response  exploiting  consistent  movements. The  horizontal  used  are  response  s e a s o n a l movement.  fitness an  to  deer's  o c c u r . The  their  These a l t e r a t i o n s  movements o f b l a c k - t a i l e d  to  be  use  p h e n o l o g i c a l changes i n the  further habitat  The range  continue u n t i l  altered.  function  observed  costs  of these h a b i t a t s  t h e s e new  altered,  selected.  use.  p a t t e r n s o f s e a s o n a l movements c o u l d  the r e l a t i v e  seasonal  as v a r i a b i l i t y  observed  the  short  t o the  may  be  evaluated,  can  f a v o u r a b l e o n e s . D e p e n d i n g on t h e l o c a t i o n  were  to t h e i r  of h a b i t a t  may  their  have been  favourable  have d i f f e r e n t  migratory  process  have t h e same n e t c o n t r i b u t i o n  These d i f f e r e n c e s  resources  by  This  h a b i t a t s . When h a b i t a t s  may  As h a b i t a t s  be  habitats.  deer  some h a b i t a t s may  fitness,  deer.  that  also  potential  in black-tailed  occupancy then  to'  can  from  one  seasonal  to the b e n e f i t s the  with  most f a v o u r a b l e the  observed  c a u s a l d i f f e r e n c e s between as  well  as  w i t h i n home r a n g e s  and  seasonal  c o s t s to habitats  p a t t e r n s of  vertical  shifts  of resident  home  deer  i n use can  and of be  161  resolved  by  The the  this  model o f h a b i t a t  model has  behaviour  of  movements  range  can  be  This  variation  in  mule d e e r  i n North  patterns  of  completely  s n o w f a l l and areas  results  i n the  with green summer  deer down  available land  migrate the  valley  bottoms  1930,  Longhurst  the  valley  into  the  summer.  In  moister  resident  winter  is  and  rains  surrounding  of  little  (Dasmann 1953,  Four  depth  and  is  to  regions  the  At  the  and  in  i f any winter.  precipitation summer  beginning  mountains but  where  green  irrigated  leave  forage  et  snowfall, and  mule d e e r Taber  and  and until  Washington  t e n d t o be  1956,  al. areas  areas  h a b i t a t s where t h e y r e m a i n  Oregon,  is  agricultural  Longhurst  these  of  instead  growth o f f o r a g e i n t h e deer  of  migratory  little  the  of the  extremes  d u r i n g the  C h a t t i n 1941,  California,  Dasmann  the  between  there  h a b i t a t s and  bottoms the  parts  snow.  and  completely  winter.  the  the  there  in  into  riparian  food.  rain  of forage  up i n t o  dry  where  senescence  the  of due  t h i s p a t t e r n of  migrate  in  The  largely  o c c u r r i n g mostly  not  After  away from  to  do  (McLean  1952).  i s low,  forage occurring  area.  i t s geographical  climatic  are  south-west  ground water  d r y i n g and  is  these  populations  from  study  w i t h i n the g e o g r a p h i c a l range  Among  dry  accounting f o r  availability  of both  movements  precipitation  away  the  t h e amount o f r a i n and  America.  the  just  throughout  form  snowpack e x i s t  In  in  seasonal  i n the  on  resident  than  availability  seasonal  populations.  deer deer  t o the  food  r e g i o n s based of the  move  mule  attributed  duration  In  of  of p r e c i p i t a t i o n  climatic  application  black-tailed  seasonal  effects  broader  selection.  Brown  mainly 1961,  162  Lauckhart  1948).  The  food  supply  does not  have t h e e x t r e m e s p a t i a l  quality  as  region  deer  (Dasmann  occurs  in  preponderance  hut  shifts  of r e s i d e n t deer  this  few  occurs  within their  are not  more s e a s o n a l l y a t t r a c t i v e third  hiqh s n o w f a l l a r e a s . In  are  a r e met  The  and  their  range  migratory  whole  deer.  year  and  i n the  summer. These c o n d i t i o n s o c c u r i n  like  Colorado,  Washington downward  and  the  in  by snow. I n  dries  out  elevations  becomes  by  e l e v a t i o n s . Deer i n t h i s c l i m a t i c  1952,  1934,  1966,  by  throughout  elevations  but  snowpacks. The  low rain  region  this the  elevations a l l o w s green  e l e v a t i o n s . In winter,  relative  the  with  are forage  snow c o v e r s  upward  elevations  in  the  high  are  Longhurst Rempel  et a l . 1977).  snowfree  areas  i s abundant deep  at  o r have  throughout  the  low  migratory  north-coastal  Here r a i n f a l l  snowpacks  is a  t o t h a t a t the  B e r t r a m and  is in  study.  year,  there  of  i s attributed  lower  forage  e t a l . 1951,  1967,  low  parts  region are p r i m a r i l y  Leopold  Loveless  fourth climatic  represented  occurs  all  Dixon  Robinette The  and  1932,  i n the  has  interior  s p r i n g t h e r e i s an  so the  more n u t r i t i o u s  and  coinciding  migration  food  summer and  and  region  winter this  which o c c u r s b e c a u s e the  m a t u r e s and  (Russell  climatic  i n h i g h e r e l e v a t i o n s and  migration  elevations  California,  this  migration  c o v e r i n g of forage  high  eastern  Oregon. . I n  altitudinal  snowfalls to  Utah,  there  mule d e e r  rainfall areas  in  range o f  the  the  The  requirements  d e e p snowpacks d u r i n g during  centres  h a b i t a t s nearby.  region winter  home  because t h e i r  heme r a n g e s f o r t h e  climatic  in  changing  quantity  of  there  seasonally  differences  the dry or  make s e a s o n a l 1953)  although  and high  shallow  summer a t  forage at high e l e v a t i o n s  163  and  the  low  available. both in  e l e v a t i o n s are Under t h e s e  resident  the  low  snowfall  shifts  in their  winter  ranges  the  winter. on  deer  the  habitat  to  as  with  a  is  useful  in  i n an  are  used  represent for and  deer.  deer  Because  bottom f o r e s t  food  (Fig.  and  mule  by  the  movements  are  same.  5).  food  and  management p r a c t i c e s deer  .  In  forest  , removal o f and  Brown  some f o r e s t e d densities Also,  than are  the  the  i s thought  1950,  these  to  1961, summer  than  is  forested  cutovers  and  night. Forest habitats  thus  p o t e n t i a l summer  types  initial  availability  Leopold  greater  of greater  the  during  subsequent h a b i t a t  herb food  area,  day  of  nutrient  1946,  i n area  during  d e e r move from  explained  the  stay  Harvesting  study  a l a r g e amount o f the  some v a l l e y  be  i n c r e a s e of forage  cutovers  have  movements o f  patterns  s h r u b and  or  more e x t e n s i v e  by  can  is  seasonal  were u n a v a i l a b l e  black-tailed  (Einarsen  make  migratory  seasonal  and  1973). I n t h e  adjacent  but  movements and  abundance o f  to deer  Hines  h a b i t a t s are  year  i n evaluating forest  canopy r e s u l t s  1968,  Forest  energy  is little  food  r e s i d e n t deer  ranges a f t e r  regions  seasonal  of  to the  and  r a n g e h a b i t a t s have e q u a l found  The  winter  climatic  of  function  beneficial  Gates  the  r e a s o n s f o r them a p p e a r t o be  where t h e r e  forest  throughout  summer r a n g e s . The  concept  regard  types  c o m p o n e n t s . The  summer h a b i t a t s t h a t  four  where  c o n d i t i o n s deer p o p u l a t i o n s  home r a n g e c e n t r e s .  their  the  predation  be  areas  Seasonal Habitats The  use  migratory  habitats  s e l e c t i o n model. A l t h o u g h t h e  different,  4.5.  only  climatic  They r e t u r n t o t h e  snowfalls in  and  the  foraging  abundance i n h i g h ( F i g . 5) ,  the  area  elevation  benefits  to  164  deer  of  logging  those  enjoyed  snowfall.  by d e e r  probably  since food  t h e study  area  are l i k e l y  i n regions further  In the study  elevations extent,  in  area  n o t a s g r e a t as  south  or  removal o f t h e f o r e s t  does n o t a f f e c t  c a n be p r o v i d e d  deer  with  canopy  lower at high  abundance t o any g r e a t  by e i t h e r f o r e s t e d o r  logged  habitats. On  summer  forested will  or cutover  depend  logging first  deer  with  results  practices cover.  would  patches  a r e only intended would  the to  high  deer's  1976).  of  after  the  Although  forest  way t o p r o v i d e  leave  patches  to  i s  initially  removed,  cover  i n the  of old-growth.  provide  cover,  deer  low  summer  Since  these  p o p u l a t i o n s d e p e n d i n g upon t h e  season  benefit or in  in  the  study  area,  replacement  deer  essentially  population. results  ranges. in  the  Logging  e x i s t at  o f f o r e s t s by  i n the r e d i s t r i b u t i o n elevation  forests  i n t h e p r o v i s i o n o f summer r a n g e s  adjacent  P r o v i s i o n of t h i s  in  which  beneficial  However, s i n c e e x t e n s i v e summer r a n g e s a l r e a d y  necessarily  some  non-merchantable  either  of  increase  deer  which would p r o v i d e t h e  a t low e l e v a t i o n s may o n l y r e s u l t  winter  by  other h a b i t a t  elevation f o r e s t s could  elevations  the  either  i n the l a c k or r e d u c t i o n of cover f o r  cutovers  to  habitats  e v a l u a t i o n i s made. I n summer, l o g g i n g may a p p e a r deer.  by  suffice.  Logging constrain  the  c o u l d be employed  ranges  stands  to  these  of  Eastman  One o b v i o u s  be  c a n be p r o v i d e d  t h e use o f  and  15 t o 20 y e a r s  harvesting  food  satisfaction  (Bunnell  usually  where  areas,  upon  requirements  the  ranges,  low  habitat  may  encourage  an  p r o p o r t i o n o f t h e r e s i d e n t component, b u t n o t  i n an i n c r e a s e o f t h e t o t a l  deer  population.  165  In winter, f o r e s t detrimental Robinson of  this  effect  f o r e s t type habitat.  will  that Some  9)  and  were used  in  these  population which d e e r  depend  population  digestible  incidentally  types  during winter  should not  be  forested  plant  with  (Fig.  9) and r e d u c e d  ( F i g . 25). Logging  the  of  of a v a i l a b l e  snow d e p t h s  forested  Robinson  logging  of these f o r e s t e d  the  p o p u l a t i o n . Edwards  winter  in  small  (19 56)  a die-off  of forested  ranges  extensively  deer  periods  shrub  and  conifer  digestible  to  logged  patches  of  suggested  by b l a c k - t a i l e d  by R o b i n s o n  that  of b l a c k - t a i l e d  black-tailed  He  noted  deer The  deer  be d e t r i m e n t a l t o  that  old-growth  deep snow  value  was  was  and r e c o g n i z e d t h e of  apparent  Sayward F o r e s t where o n l y d e e r remaining  deer i n  were i n o l d - g r o w t h  would  (1956) o b s e r v e d  ranges.  (1956).  in  i n deep s n o w f a l l r e g i o n s  winter ranges  winter  dry matter  d u r i n g p e r i o d s o f deep snow  winter ranges  forests.  importance  the  or snow-free  d u r i n g p e r i o d s w i t h deep snowpacks, d e e r  factor  to  cutovers as winter f o r a g i n g areas  deep s n o w f a l l r e g i o n s was o b s e r v e d  a  (Fig.  a s s o c i a t i o n s a t low e l e v a t i o n s p r o v i d e  greater densities  Dse  deer  deer  ( F i g . 9) .  deer  that  winter  associations  for  detrimental  would be g r e a t e s t once t h e y had r e a c h e d  winter.  , the  t h e c u t o v e r s c r e a t e d may p r o v i d e f o r a g i n g a r e a s  w i n t e r . The v a l u e o f t h e s e  Other  magnitude  of other  plant  a  1956,  of the winter  d r y matter  c o u l d use d u r i n g low snow d e p t h  serai stages  The  and the a v a i l a b i l i t y forested  have  (Edwards  1978).  upon t h e s e v e r i t y  elevation,  available  and  deer  1974 and 1975, Bloom  low  little  forest  the  i s logged,  provided  in  on  1956, J o n e s effect  h a r v e s t i n g a t low e l e v a t i o n s may  forested in  the  inhabiting the  survived  the  winter  166  (Edwards habitat and  1956).  Guiguet  forest  (1965) and  deer  In taken that of  deer  was  further  Alaska Dept. F i s h  priority  that  were  used  are s i m i l a r  deer  in  deer  habitat  habitat  population.  winter  range"  reguirements  h a b i t a t s may  be  reserves  with  does n o t  follow  the deer The  greater shallow  total  densities winter  range.  to  consideration  severe  of  Game  (1973).  about  Jones  the  types  during winter. used  "critical  is  not  by  The  radio-  be  should the  the deer  p o p u l a t i o n , and  assumption  is  preservation  component  deer  and  of  other  winter  (Mautz  winter range"  the  seasonal  with adequate  energy  1978). I t  i s preserved  maintained.  winter may  deer  ranges  may  have  p r o v i d e the d a y s o f use.  plant an  can  be  winter energy not j u s t  and  greater  During  have  associations area  overestimated deer  population with  as  be g i v e n t o t h e c o n t r i b u t i o n  meeting  has  necessarily true since  winter ranges  evaluating  winter range"  means  isolated  mild winter ranges  towards  habitat.  and  those  can s u r v i v e t h e  some f o r e s t e d In  Cowan  i n p r o v i d i n g deer  of severe  number  than  by  management. The  an  winter ranges  snow d e p t h s  acknowledged  deer  black-tailed  population w i l l  , mild  by  i f a l l "critical  because even though densities  This  of  they  that  importance  winter  winter range"  i s not  important  which  as  study.  p r e s e r v a t i o n of " c r i t i c a l the  forest  (1978) were more s p e c i f i c  identified  in this  of old-growth  recent years preservation of  "critical  then  Bloom  habitat  h a b i t a t s they tagged  necessity  for black-tailed  (1974,1975) and of  The  nutrient  the d e n s i t y of  periods of  greater used  as  of the  food severe  winter  range, habitat  requirements deer  a  in  of  that  167  Failure  to  provide  d e t r i m e n t a l t o the deer winter for  range.  deer  winter  only severe  deer  the  even p e r m i t increasing ranges  to  winter they  entire  reduce  them t o their  are  a  provided along  with  to severe  deer  winter  then  range  Severe  1975)  and  Douglas-fir  the  more  study  severe  m i l d e r and snow d e p t h s fir  association  Amabilis from  severe winter ranges winter ranges  than  d i d the severe  that severe  lichens winter  are  severe  of mild  winter  shallow loss  snow  o r i t may  I f mild  thus winter then  entire  p o p u l a t i o n could cause  an  t o a degradation of the  are  Deer used  this  periods  in  into  Salal  by  Jones  primarily  association  more  ranges.  left  type  -  the S a l a l Western  and  better  Stevenson  difference  in  -  -  mainly was  Douglas-  Hemlock  S i n c e deer  and moved  suggests  that  a t these  times  (1978)  a b u n d a n t on m i l d w i n t e r This  (1974  w i n t e r . When w e a t h e r  requirements  winter ranges.  winter  i n the S a l a l  during mild periods i t  met t h e i r  black-tailed  m i l d and s e v e r e  are described area  for  W e s t e r n Hemlock a s s o c i a t i o n s .  mild  their  winter ranges f o r the  s h a l l o w e r , deer  and moved  Fir -  i s provided  severe winter ranges,  h a b i t a t i s t o be p r o v i d e d  association.  the  severe  carrying capacity.  winter ranges  in  j u s t as  balance,  survival.  i t must i n c l u d e a r e a s o f b o t h  range.  during  energy  o f t h e f o r a g e and l e a d  winter range's  If  o f energy  positive  w i n t e r . T h i s c o n c e n t r a t i o n of t h e deer over-utilization  habitat  during periods of rate  be  t o provide  would be on  chances of o v e r - w i n t e r  not  may  winter. Provision  their  secure  would be r e s t r i c t e d  severe  range,  w i n t e r range  which would be a v a i l a b l e  allow  deer  for  winter  p o p u l a t i o n as f a i l u r e  d u r i n g each  diet  ranges may  then  If  mild  has  ranges  lichen  shown  than  on  abundance  168  combined area  w i t h g r e a t e r amounts o f s h r u b s  of  mild  winter  ranges,  s u b s t a n t i a l food  source  Because o f t h e i r  potential  may  be  just  forest  o f the deer  maintained  British that  of  deer.  Columbia  "long  valley  winter  mild  ranges.  winter  winter ranges  ranges  corridors  ranges  t o the over-  s u b a l p i n e summer  and W i l d l i f e  with  summer  -  plans  land  Results of this corridors"  and  migrations should summer  snowfalls.  mature and low  downward  winter  (1976)  who  habitat  recommend  extending  established The  to  provision are  by t h e  from  connect  of  these  implemented  i t s input  into  by  forest  1977) .  value  that  the  concept  in facilitating  be r e a s s e s s e d . M i g r a t i o n  t o w i n t e r home r a n g e s  of  "deer  black-tailed  of  some  deer  o c c u r r e d soon a f t e r  deer from  the f i r s t  These s n o w f a l l s were e p h e m e r a l and n o t deep enough t o  deer  movement. S e v e r a l t i m e s  the w i n t e r deer  moved t o t h e i r  and r e t u r n e d t o t h e i r the  be  through  study suggest  their  of  ranges  corridors"  ranges.  Service  (Young  Branch  deer  i n t h e management o f d e e r  harvesting  strips  c o r r i d o r s a r e a l s o suggested  firebreak  B. C. F o r e s t  prevent  continuous  i n order to f a c i l i t a t e  These  Fish  term  that  bottoms t o h e i g h t s o f  winter  when  severe  a more  population.  between  winter ranges  migration  line  with  on  as severe  (1975) p r o p o s e d  be  elevation  their  provide deer  Migration Corridors Jones  the  that  and g r e a t e r  as f o r a g i n g a r e a s ,  as i m p o r t a n t  winter s u r v i v a l  4.6.  than  would  and c o n i f e r s  snow  melted.  during the i n i t i a l  w i n t e r home r a n g e s  summer home r a n g e s Great  mobility  part of  below t h e snow  a few  days  later  i s also evident i n that  169  migratory Given with  movements were a c c o m p l i s h e d  the  sensitivity  which  migration  involved, tailed  it  deer  is  corridors  can  that  and  reported  i n mule d e e r  (Russell  1932,  and  1971).  been  Longhurst  Although  reported  and  1971),  no  Olsen  Robinette  1960,  numbers o f b l a c k by  deep snow  i f  no  and  forested  where m i g r a t i o n s o f  of  their  et  Walden  that  deer  and  are unnecessary.  I f cover f o r  other  species i s desired  or s t a g g e r e d f o r e s t e d  placement  and  Forested as  decreases  function  of c o r r i d o r s  forested  areas a l l o t t e d were  is  to  with  horizontally  Verme and  Ozoga  deer  caused  their  are  sensitivity  deer  by  even i n  common.  or  The  t o snow routes  habitat  by  the  for  strips. proposed  by  increasing mild  purpose  deployed  by  appropriate  Jones  (1975)  to  use  of  p o t e n t i a l f o r winter  provide  for this  1970,  t h e n i t c o u l d be p r o v i d e d  also The  1956,  km  b l o c k s , or  were  mild winter range.  corridors  80  1951,  Edwards  c o r r i d o r s as m i g r a t o r y  enhancement o f r e s i d u a l corridors  ranges  1952,  mule  migrating  segmented  Wallmo  winter  have been o b s e r v e d ,  over  p r o v i s i o n of f o r e s t e d  wildlife  of  been  Leopold et a l .  al.  summer r a n g e s  black-tailed  has  L e o p o l d e t - a l . . 1951,  i n s t a n c e s o f mass d i e - o f f s  populations  they  speed  distances  snowfalls  1947,  Cumming and  on  if  , the  short  d i e - o f f s o f u n g u l a t e s on  entrapment  these  early  (Severinghaus  e t a l . 1952,  Klein  function  hours.  existed. and  suggest  12  deer  winter ranges,  weather  mobility  the  substantial  to  have  than  trapped a t high e l e v a t i o n s  t o move t o t h e i r  Sensitivity  Gill  less  black-tailed  occur,  unlikely  would be  t h u s be u n a b l e  t o snow by  in  and  altitude. winter  would  be  bordered  range  more  I f the then  effective  severe  winter  170  ranges.  This  orientation  accessible  to  longer  i n winter  time  corridors  a greater  between h i g h  would  proportion  and and  be  make of the  mild  elevations.  ranges  deer p o p u l a t i o n  more b e n e f i c i a l  low  winter  than  the  for a  present  171  5.  Literature Cited ;  Adams, C.C. 1919. Migration as-a factor i n evolution: i t s e c o l o g i c a l d y n a m i c s . I I . Am. N a t . 53: 55-78. A l a s k a Dept. habitat.  Fish and Game. A l a s k a Dept. F i s h  1973. Alaskans wildlife and Game. Pages 9-11.  and  A l e x a n d e r , J . E . 1973. S e a s o n a l movements o f e l k . A l a s k a Dept. Fish And Game. J u n e a u , A l a s k a . F i n . S e p t . F e d . A i d W i l d l . B e s t . P r o j . W - 1 7 - 3 and W-17-4 J o b 13.1 B. 37 p. A l e x a n d e r , B.D. 1974. The e v o l u t i o n Eev. E c o l . S y s t . 5:325-383.  of s o c i a l  behavior.  Annu.  Ammann, A.P., B.L. Cowan, C.L. M o t h e r s h e a d , and B.R. Baumgardt. 1973. Dry m a t t e r and e n e r g y intake i n relation to d i g e s t i b i l i t y i n w h i t e - t a i l e d deer. J . W i l d l . Manage. 37: 195-201. Ashcraft, G.C., J r . 1961. Deer movements o f t h e McCloud herds. C a l i f . F i s h Game 47: 145-152.  Flat  B a i l e , C A . And J.M. F o r b e s . 1974. C o n t r o l o f f e e d i n t a k e and regulation o f energy b a l a n c e i n ruminants. P h y s i o l . Eev. 54: 160-214. Bell,  M.A.M. 1971. F o r e s t ecology. Pp, 200-287Forestry handbook for British Columbia . The F o r e s t Club, U n i v e r s i t y o f B r i t i s h C o l u m b i a . 815 p.  B e r g e r u d , A.T. 1974. The r o l e o f the environment i n the aggregation, movement, and disturbance behaviour of caribou. Pp. 552-584 I n : V. G e i s t a n d F . W a l t h e r , e d s . The b e h a v i o u r o f u n g u l a t e s and i t s r e l a t i o n t o management, V o l . 2. IUCN P u b l . No. 2 4 . 940 p. Bertram, R.C. and E.D. Eempel. Kings deer herd. C a l i f . Fish  1977. M i g r a t i o n o f t h e N o r t h Game 63: 157-179.  B l a i r , W.F. 1953. P o p u l a t i o n d y n a m i c s of s m a l l mammals. Adv. G e n e t . 5:1-41. Blaxter, K.L. Hutchinson Blood,  rodents  1962. The e n e r g y metabolism And C o . L t d . , L o n d o n . 329 p.  of  and  other  ruminants.  D.A. 1963. Some a s p e c t s o f b e h a v i o r o f a b i g h o r n Can. F i e l d - N a t . 77: 77-94.  herd.  Bloom, A.M. 1978. S i t k a b l a c k - t a i l e d d e e r w i n t e r range i n t h e K a d a s h a n Bay a r e a , s o u t h e a s t A l a s k a . J . W i l d l . Manage. 42: 108-112.  172  B o u c k h o u t , L.W. 19 72. The b e h a v i o u r o f mule d e e r ( O d o c o i l e u s hemionus hemionus R a f i n e s q u e ) i n w i n t e r i n r e l a t i o n t o t h e s o c i a l and p h y s i c a l e n v i r o n m e n t . M.Sc. T h e s i s , University of C a l g a r y . 124p. British Columbia Fish and W i l d l i f e B r a n c h . 1976. G u i d e l i n e s for the protection of w i l d l i f e habitat i n coastal British Columbia. B.C. F i s h and W i l d l . B r . 14 p. T y p e w r i t t e n . Brooke, R.C., E.B. P e t e r s o n , and s u b a l p i n e m o u n t a i n hemlock zone. 147-349. Brown,  V.J. Krajina. 1970. The E c o l . W e s t e r n N. Am. 2:  E . S . 1961. The b l a c k - t a i l e d d e e r o f w e s t e r n Washington. W a s h i n g t o n S t a t e Game D e p t . B i o l . B u l l . 13: 1-124.  B u n n e l l , F . L . And D.S. Eastman. 1976. Effects of forest management practices on wildlife i n forests of British C o l u m b i a . Pp. 631-689 i n P r o c . D i v . I . XVI IUFRO World C o n g r e s s , O s l o , Norway. Burt,  W.H. 1943. Territoriality and home r a n g e a p p l i e d t o mammals. J . Mammal. 24:346-352.  c o n c e p t s as  B y f o r d , J ^ L . 1969. Movement r e s p o n s e s o f - w h i t e - t a i l e d d e e r t o changing food supplies;. P r o c . Annu. Conf, S o u t h e a s t A s s o c . Game F i s h Comm. 23: 63-78. Cameron, R.D., R.G. White, and J.R. L u i c k . 1975. The a c c u m u l a t i o n o f water i n r e i n d e e r d u r i n g w i n t e r . . Pp. 374378. I n : J.R. L u i c k , P.C. L e n t , D.R. K l e i n , and R.D. White (eds.). Proc. 1 s t I n t l . C a r i b o u R e i n d e e r Symp. B i o l . Pap. U n i v . A l a s k a S p e c . Rept, 1: 1-551. Capp,  J.C. 1968. Bighorn sheep, e l k , mule deer range relationships. Dep. F i s h e r y and W i l d l . B i o l . Colorado State U n i v e r s i t y . 75 p.  C l o v e r , M..R. 1956. 199-201.  Single  gate  deer t r a p .  C o b l e n t z , B.E. 1970. Food h a b i t s W i l d l . Manage. 34: 535-539.  Calif.  Fish  o f George Reserve  Game 42:  deer.  J.  Cohen, D. 1967. O p t i m i z a t i o n o f s e a s o n a l m i g r a t o r y b e h a v i o u r . Am. N a t . . 1 0 1 : 5-17. Cook,  G.W. 1972. Comparative nutritive values of forbs, grasses, and s h r u b s . Pp. 303-310 I n : C M . M c K e l l , J . P . B l a i s d e l l , and J.R. G o o d i n ( e d s . ) , W i l d l a n d s h r u b s their biology and u t i l i z a t i o n . USDA F o r e s t S e r v i c e Gen. T e c h . R e p t . INT-1.  C o v i c h , A.P. 1976.. A n a l y z i n g s h a p e s o f ecological and economic t h e o r i e s . 7:735-757.  foraging areas: some Annu. Rev. E c o l . S y s t .  173  Cowan, I.McT. 1945. The e c o l o g i c a l r e l a t i o n s h i p s o f t h e f o o d of the Columbian black-tailed d e e r , O d o c o i l e u s hemionus columbianus (Bichardson) , i n t h e c o a s t forest region of s o u t h e r n Vancouver I s l a n d , B r i t i s h Columbia. E c o l . Monogr. 15: 110-139. Cowan, I . McT. 1956. What and where a r e t h e mule and b l a c k t a i l e d deer? Pp. 335-359 I n : W.P. T a y l o r ( e d . ) . T h e d e e r of North America. The S t a c k p o l e C o . , H a r r i s b u r g , P a . And t h e W i l d l i f e Management I n s t i t u t e , W a s h i n g t o n , D.C. 668.p. Cowan, I . McT. and C . J . G u i g u e t . 1965. The mammals o f B r i t i s h Columbia. B.C. P r o v . Mus. Handb..11. 414 p. C r a i g h e a d , J . J . , F.C. C r a i g h e a d J r . , E . L . B u f f , and B.W. O'Gara. 1973. Home r a n g e s and a c t i v i t y p a t t e r n s of non-migratory elk of t h e Madison drainage herd a s determined by telemetry. W i l d l . Monogr. 33:1-50. Crampton, E.W. Nutrition.  And W.H.  L.E. H a r r i s . , 1969. Applied Animal Freeman And Co., San F r a n c i s c o . 753 p.  Crook, J.H. 1970. The s o c i o - e c o l o g y o f p r i m a t e s . Pp.103-166. I n : J.H. C r o o k (ed.). Social behaviour i n b i r d s and mammals. Academic P r e s s , London. 492 p. Crook,  J.H., J-E. Ellis, Mammalian s o c i a l s y s t e m s : Behav. 24:261-274.  and J.D. G o s s - C u s t a r d . s t r u c t u r e and f u n c t i o n .  1976. Anim.  Cumming, H.G. and F.A. Walden. 1970. The w h i t e - t a i l e d d e e r i n Ontario. O n t . Dep. L a n d s and F o r e s t s , F i s h and W i l d l . B r . 25 p. Dalke, P.D., E. D. Beeman, F . J . K i n d e l l , E . J . B o b e l and T. R. Williams. 1965. S e a s o n a l movements o f e l k i n t h e Selway E i v e r Drainage. J . W i l d l . Manage. 29: 333-338. Dalke, P.D. and P.E. Sime. 1938. Home and s e a s o n a l r a n g e s o f the e a s t e r n c o t t o n t a i l i n Connecticut. Trans. N. Am. W i l d l . C o n f . 3: 659-669. Dasmann, E.F. 1953. Factors influencing movement o f nonmigratory deer. P r o c . Annu. C o n f . W e s t e r n A s s n . Game Fish Comm. 33: 112-116. Dasmann, E . F . And E.D. T a b e r . 1956. B e h a v i o r o f C o l u m b i a n b l a c k - t a i l e d d e e r with r e f e r e n c e t o p o p u l a t i o n e c o l o g y . J . Mammal. 37: 143-164. D a v i e s , N.B. 1977. P r e y s e l e c t i o n and t h e s e a r c h strategy of the. spotted flycatcher ( Muscicapa striata ) : a field s t u d y on o p t i m a l f o r a g i n g . Anim. Behav. 25: 1016-1033. D a v i s , D.E., J . T . Emlen J r . , a n d A.W. S t o k e s . 1948. S t u d i e s o f home r a n g e i n t h e brown r a t . J . Mammal. 29: 207-225.  174  Dealy, J.E. 1959. The influences of logging Columbian b l a c k - t a i l e d deer in the Blue Oregon. M.Sc. Thesis. Oregon S t a t e C o l l e g e . Dixon, J . 1934. A study of the l i f e mule d e e r i n C a l i f o r n i a . Calif. Edie,  practices Biver area 65 p.  on of  h i s t o r y and f o o d h a b i t s F i s h Game 20: 181-282.  of  A. and A. H a r e s t a d . 1971. Investigations of elk other wildlife on V a n c o u v e r I s l a n d , May-Sept..1971. F i s h and W i l d l . B r . , Nanaimo. 37 p. Typewritten.  Edwards, R.Y. mountains 168.  1956. Snow d e p t h s and of western Canada.  u n g u l a t e abundance i n J . W i l d l . Manage. 20:  Edwards, R.Y. And R.W. Ritcey. 1956. herd. J . Mammal. 37: 486-494. E i n a r s e n , A.S. 1946. W i l d l . Manage. 10: Eisenberg, J.F. Handb. Z o o l .  Management 54-59.  1966.. The 10: 1-92.  The  of  social  migrations  black-tailed organization  of  the 159-  a moose  deer. of  and B.C.  J.  mammals.  E s t e s , R.D. 1974. S o c i a l o r g a n i z a t i o n of the A f r i c a n Bovidae. Pp. 166-205. In : V. Geist and F. W a l t h e r ( e d s . ) . The b e h a v i o u r of u n g u l a t e s and i t s r e l a t i o n t o management. IUCN P u b l . No. 24. V o l . I . Morges, S w i t z e r l a n d . 511 p. F i t z h a r r i s , B.B. 1975. Snow a c c u m u l a t i o n and d e p o s i t i o n on a west coast mid-latitude mountain.. Ph.D. Thesis. U n i v e r s i t y of B r i t i s h C o l u m b i a . 367 pp. G a t e s , B.R. 1968. Deer f o o d p r o d u c t i o n of the c o a s t f o r e s t . M.Sc. Thesis. Columbia . 105 p. Geist, V. ecology  1974. On t h e i n ungulates.  i n c e r t a i n serai stages U n i v e r s i t y of British  r e l a t i o n s h i p of s o c i a l e v o l u t i o n Am. Z o o l . 14: 205-220.  G i l b e r t , N.E. 1973. Biometrical P r e s s , O x f o r d . . 1 2 5 p.  interpretation.  and  Clarendon  Gilbert, P.F., O.C. Wallmo, and R.B. G i l l . 1970. Effect snow d e p t h on mule deer in Middle Park, Colorado. W i l d l . Manage. 34: 15-23. G r u e l l , G. 1958. R e s u l t s o f f o u r y e a r s o f t r a p p i n g and deer in northeastern Nevada. P r o c . Annu. C o n f . A s s o c . Game F i s h Comm. 38: 179-183.  of J.  tagging Western  H a r d i s o n , W.A. 1959. Evaluating the nutritive quality of forage on the b a s i s of energy - a review. J . Dairy S c i . 42: 489-500.  175  Harestad, body  A.S. And F.L. B u n n e l l . (in press). weight - a r e e v a l u a t i o n . Ecology.  Home  range  and  Hebert, D.M. 1973. A l t i t u d i n a l m i g r a t i o n as a f a c t o r i n the n u t r i t i o n of b i g h o r n sheep. Ph.D. T h e s i s . , University of B r i t i s h C o l u m b i a . 357 p . Hepburn, R.L. 1959. Effects of snow c o v e r on m o b i l i t y and local distribution of deer in Algonquin Park. M.Sc. Thesis. University of Toronto. 55 p. O r i g i n a l n o t s e e n . C i t e d f r o m : Eastman, D.S. 1978. H a b i t a t s e l e c t i o n and use in winter by moose i n s u b - b o r e a l f o r e s t s o f n o r t h c e n t r a l B r i t i s h Columbia, and relationships to forestry. Ph.D. T h e s i s . U n i v e r s i t y o f B r i t i s h C o l u m b i a . 554 p. Hines, W.W. 1973. B l a c k - t a i l e d d e e r p o p u l a t i o n s and D o u g l a s f i r r e f o r e s t a t i o n i n the Tillamook Burn, Oregon. Oreg. Game Comm. Res. R e p t . 3: 1-59. Hirth, D.H. relation  1977. Social behavior of w h i t e - t a i l e d to habitat. W i l d l . Monogr. 53: 1-55.  Hoskinson, R.L. And L.D. Mech. migration and i t s role in Manage. 40: 429-441.  deer i n  1976. White-tailed wolf predation. J.  deer Wildl.  H o s l e y , N.W. 1956. Management of white-tailed deer in its environment. Pp. 187-259 I n : W.P. Taylor (ed.). The d e e r of North America. S t a c k p o l e Co. H a r r i s b u r g , Pa. and t h e W i l d l i f e Management I n s t i t u t e , W a s h i n g t o n , D . C . 6 6 8 p. Jacobsen, N.L.K. 1973. Physiology, behavior, and transactions of w h i t e - t a i l e d deer. Ph.D. T h e s i s . U n i v e r s i t y , I t h a c a . 245 p. Jarman, P . J . relation  1974. The social to t h e i r ecology.  organization B e h a v i o u r 48:  Jones, G.M. 1972. C h e m i c a l f a c t o r s and t h e i r i n t a k e r e g u l a t i o n i n ruminants : a review. S c i . 52: 207-239.  thermal Cornell  of antelope 213-267.  in  r e l a t i o n to feed Can. J. Anim.  Jones, G.W. 1974. I n f l u e n c e o f f o r e s t d e v e l o p m e n t on b l a c k t a i l e d d e e r w i n t e r r a n g e on V a n c o u v e r I s l a n d . Pp. 139-148. In H.C. Black, ed. Proc. Symp. Wildlife and forest management in the Pacific Northwest. Oregon S t . U n i v . , C o r v a l l i s , O r e g o n . 236 p. J o n e s , G.W. 1975. Aspects of the winter ecology of blacktailed deer ( O d o c o i l e u s hemionus c o l u m b i a n u s R i c h a r d s o n ) on n o r t h e r n V a n c o u v e r I s l a n d . M.Sc. T h e s i s . University of B r i t i s h Columbia . 78 p. K e l s a l l , J.P. 1969. S t r u c t u r a l a d a p t a t i o n s o f moose f o r snow. J . Mammal. 50: 302-310.  and  deer  176  Klein, D.R. 1965. Ecology Monogr. 3 5 : 259-284.  of  deer  range  i n Alaska.  Ecol.  K l e i n , D.R. 1970. Food s e l e c t i o n by N o r t h American d e e r and their response to over-utilization of p r e f e r r e d plant species. Pp. 25-46 I n : A. Watson (ed.). Animal populations i n r e l a t i o n t o t h e i r food resources. B l a c k w e l l S c i e n t i f i c P u b l . , O x f o r d a n d E d i n b u r g h . 477 p. K l e i n , D.R. And S.T. O l s e n . 1960. N a t u r a l m o r t a l i t y p a t t e r n s o f deer i n s o u t h e a s t A l a s k a . J . W i l d l . Manage. 24: 80-88. K n o w l t o n , F . F . 1960. F o o d h a b i t s , movements, and p o p u l a t i o n s of moose i n t h e G r a v e l l y M o u n t a i n s , Montana. J. Wildl. Manage. 24: 162-170. K o w a l s k i , C . J . 1970. The p e r f o r m a n c e o f some r o u g h tests f o r bivariate normality before and after coordinate transformations to normality. T e c h n o m e t r i c s 12: 517-544. K r a j i n a , V . J . 1965. B r i t i s h Columbia  B i o g e o c l i m a t i c z o n e s and c l a s s i f i c a t i o n o f . E c o l . W e s t e r n N. Am. 1: 1-17.  K r u m l i k , G . J . 1979. C o m p a r a t i v e s t u d y o f n u t r i e n t cycling i n the subalpine Mountain Hemlock Zone o f B r i t i s h Columbia. Ph.D. T h e s i s . O n i v . Of B r i t i s h C o l u m b i a . 191 p. K u c e r a , T . E . 1978.. S o c i a l b e h a v i o r a n d b r e e d i n g d e s e r t mule d e e r . J . Mammal. 59: 463-476.  systems o f t h e  L a u c k h a r t , J . B . 1948. B l a c k - t a i l e d d e e r i n w e s t e r n Washington. Proc. Annu. C o n f . W e s t e r n A s s o c . Game F i s h Comm. 28: 15316 1. Leege, T.A. And W.O. Hickey. 1977. Elk-snow-habitat r e l a t i o n s h i p s i n the Pete King Drainage, Idaho. I d a h o Dep. F i s h Game W i l d l . B u l l . 6: 1-23. Leopold, A.S. 1950. Deer i n r e l a t i o n t o plant succession. T r a n s . N. Am. W i l d l . C o n f . 15: 571-580. L e o p o l d , A.S., T. R i n e y , £ . M c C a i n , and L . T e v i s J r . 1951. The Jawbone d e e r h e r d . C a l i f . Dep. F i s h Game Game B u l l . 4: 1139. LeResche, Can. List,  R.E. 1974. Moose m i g r a t i o n s i n N o r t h (Que.) 101: 393-415.  America.  R . J . 1966. S m i t h s o n i a n m e t e o r o l o g i c a l t a b l e s . M i s c . C o l l e c t . 114: 509.  L o n g h u r s t , W.M. And J . E . C h a t t i n . F i s h Game 27: 2-12.  1941.  Smithson.  The b u r r o d e e r .  L o n g h u r s t , W.M., A.S- L e o p o l d , and R.F. survey of California deer herds,  Nat.  Calif.  Dasmann. 1952. A their r a n g e s , and  177  management problems. 1-136.  C a l i f . Dep. F i s h Game Game  Bull.  6:  L o v e l e s s , C.W. 1967. E c o l o g i c a l c h a r a c t e r i s t i c s o f a mule deer w i n t e r range. C o l o . D i v . Game F i s h P a r k s Tech. P u b l . 20: 1-124. MacArthur, R.H. and E.B. P i a n k a . 1966. On o p t i m a l patchy environment. Am. Nat. 100: 603-609.  use o f a  Mackie, E . J . 1970. Eange e c o l o g y and r e l a t i o n s of mule d e e r , e l k , and c a t t l e i n t h e M i s s o u r i B i v e r b r e a k s , Montana. W i l d l . Monogr. 120: 1-79.. Mautz, W.W. 1978. S l e d d i n g on a bushy h i l l s i d e i n d e e r . W i l d l . Soc. B u l l . 6: 88-90.  : the f a t c y c l e  M c C u l l o u g h , D.E. 1964. R e l a t i o n s h i p s o f weather t o migratory movements of b l a c k - t a i l e d deer. E c o l o g y 45: 249-256. McLean, D.D. 1930. The Burro deer i n C a l i f o r n i a . Game 16: 119-120.  Calif.  Fish  McNab, B.K. 1963. B i o e n e r g e t i c s and t h e d e t e r m i n a t i o n o f home range s i z e . Am. Nat. 97: 133-140. Metzgar, L.H. 1967. An e x p e r i m e n t a l comparison o f s c r e e c h o w l p r e d a t i o n on r e s i d e n t and t r a n s i e n t white-^f ooted mice (Peromyscus l e u c o p u s ) . J . Mammal. 48: 387-391. Miller, F.L. 1970. D i s t r i b u t i o n p a t t e r n s of b l a c k - t a i l e d deer (Odocoileus hemionus columbianus) i n relation to environment. J . Mammal. 5 1 : 248-260. Moir,  B.C. 1976. Movement and range use p a t t e r n s of mule deer i n t h e P a h s i m e r o i V a l l e y , Idaho. M.Sc. T h e s i s . University of Idaho. 89 p.  Moir,  R.J. , 1961. A note on t h e r e l a t i o n s h i p between t h e d i g e s t i b l e dry matter and t h e d i g e s t i b l e energy c o n t e n t o f ruminant a n a l y s e s . Aust. J . Exp. A g r i c . Anim. Husb. 1: 2426.  Morgan, C L . 351 p.  1896. H a b i t and i n s t i n c t .  E. A r n o l d Co., London.  N e f t , D.S. 1966. S t a t i s t i c a l a n a l y s i s f o r a r e a l d i s t r i b u t i o n s . Regional Science Research Institute, Pennsylvania. Monograph S e r i e s Number 2. 179 p. Nordon, H.C., I . McT. Cowan, and A . J . Wood. 1970. The f e e d i n t a k e and heat p r o d u c t i o n o f young b l a c k - t a i l e d deer (Odo c o i l eus hemionus Columbian u s ) . Can. J . Z o o l . 48: 275282.  178  Oldemeyer, J.L. Can. (Que.)  1974. N u t r i t i v e v a l u e o f moose f o r a g e . 101: 217-226.  O r l o c i , L. 1964. V e g e t a t i o n a l and the ecosystems o f the c o a s t a l Thesis. O n i v e r s i t y of B r i t i s h O r r , R.T. 1970. 303 p.  Nat.  environmental v a r i a t i o n s in w e s t e r n hemlock zone. Ph.D. C o l u m b i a . 206 p.  Animals i n m i g r a t i o n .  M a c M i l l a n Co.,  London.  Person, S.J., R.G. White, and J.E. Luick. 1975. In v i t r o d i g e s t i b i l i t y o f f o r a g e s u t i l i z e d by Ranqifer tarandus Pp. 251-256.. I n : J.R. L u i c k , P.C. L e n t , D.R. K l e i n , and R.G. White ( e d s . ) . P r o c . 1 s t I n t l . Reindeer And Caribou Symp. B i o l . Pap. O n i v . A l a s k a Spec. Rept, 1: 1-551. Phillips, R.L., W.E. Berg and D.B. Siniff. 1973. Moose movement p a t t e r n s and r a n g e use i n n o r t h w e s t e r n Minnesota. J . W i l d l . Manage. 37: 266-278. Pyke,  G.H., H.R. Pulliam, and E.L. C h a r n o v . foraging: a s e l e c t i v e r e v i e w o f t h e o r y and Rev. B i o l . 52: 137-154.  Reid,  R.L. 1968. R a t i o n s f o r m a i n t e n a n c e and p r o d u c t i o n . Pp. 190-200. I n : F.B. Golley and H. K. Buechner (eds.) A practical guide t o t h e study of the p r o d u c t i v i t y of l a r g e h e r b i v o r e s . IBP Handbook 7. B l a c k w e l l S c i . Publ. Oxford. 308 p.  Rittenhouse, L.R., C.I. Streeter, Estimating digestible energy organic matter in diets of Manage. 24: 73-75. Robinette, Otah.  1977. tests. i  Optimal Quart.  and D.C. Clanton. 1971. from digestible dry and grazing c a t t l e . . J . Range.  W.L. 1966. Mule d e e r home r a n g e J . W i l d l . Manage. 30: 335-349.  and  dispersal  in  Robinette, W.L., 0. J u l a n d e r , J . S . G a s h w i l e r , and J.G- S m i t h . 1952. W i n t e r m o r t a l i t y o f mule d e e r i n Utah i n r e l a t i o n t o range c o n d i t i o n . J . W i l d l . Manage. 16: 289-299. R o b i n s o n , D.J. 1956. P r e l i m i n a r y s t u d i e s upon the effect of logged over a r e a s on q u a l i t y and s i z e o f d e e r p o p u l a t i o n s on Vancouver Island. Annu. Meeting of the Northwest S e c t i o n o f t h e W i l d l i f e S o c i e t y , S e a t t l e , W a s h i n g t o n . 8 p. Rochelle, J.A. (in preparation). The r o l e of mature c o n i f e r f o r e s t s i n t h e w i n t e r n u t r i t i o n o f b l a c k - t a i l e d d e e r . Ph.D. Thesis. U n i v e r s i t y of B r i t i s h Columbia . Russell, CP. 1932.. S e a s o n a l m i g r a t i o n o f Monogr. 2: 1-46. S c h o e n e r , T.W. 1968. S i z e s of f e e d i n g E c o l o g y 49: 123-141.  mule  territories  deer. among  Ecol. birds.  179  S c h o e n e r , T.W. 1971. T h e o r y E c o l . S y s t . 2: 369-404.  of feeding  strategies.  Annu. Rev.  Seton, E.T. 1909. Life histories of northern a c c o u n t o f t h e mammals o f M a n i t o b a . V o l . I . C. Sons, New Y o r k . 673 p.  a n i m a l s . An Scribners's  Severinghaus, C.W. 1947. Relationship o f weather t o w i n t e r mortality and p o p u l a t i o n levels among deer in the Adirondack r e g i o n o f New Y o r k . Trans.. N. Am. W i l d l . C o n f . 12: 212-223. S i e g e l , S. 1956. N o n - p a r a m e t r i c s t a t i s t i c s f o r sciences. McGraw H i l l Book Co., New York.  the behavioural 312 pp.  S m i t h , I.D. 1968. The e f f e c t s o f s e r a i s u c c e s s i o n and h u n t i n g upon Vancouver Island black-tailed d e e r . . M.Sc. T h e s i s . U n i v e r s i t y of B r i t i s h Columbia . 140 p. S m i t h , I.D. 1972. A management plan f o r deer in British Columbia. B.C. F i s h and W i l d l . B r . 23 p . , T y p e w r i t t e n . Smith, I.D. and R. D a v i e s . 1973. A p r e l i m i n a r y investigation o f t h e c h a r a c t e r i s t i c s o f d e e r and e l k r a n g e i n t h e T s i t i k a R i v e r watershed, Vancouver I s l a n d . B.C. F i s h and Wildl, B r . , Nanaimo. 129 p. T y p e w r i t t e n . Smith, J.N.M. and H.P.A. Sweatman. 1974. Food searching behaviour of t i t m i c e i n patchy environments. Ecology 55: 1212-1232. Sokal, R.R. a n d F . J . R o h l f . 1969. Co., San F r a n c i s c o . 776 p .  Biometry.  W.H.  Freeman a n d  Sparrow, R.D. And P.P. S p r i n g e r . patterns of whit-tailed deer Manage. 34: 420-431.  1970. Seasonal activity i n South Dakota. J . Wildl.  S t e l f o x , J.G. 1976. Range e c o l o g y o f Rocky Mountain sheep. Can. W i l d l . S e r v . R e p t . S e r . 39. 50 p.  bighorn  Stevenson, S.K. 1978. D i s t r i b u t i o n and abundance o f a r b o r e a l l i c h e n s a n d t h e i r use a s f o r a g e by b l a c k - t a i l e d d e e r . M.Sc. Thesis. U n i v e r s i t y o f B r i t i s h Columbia . 148 p. S t r a n d g a a r d , H. 1972. The r o e d e e r ( Capreolus capreolus ) population a t Kalo and t h e f a c t o r s r e g u l a t i n g i t s s i z e . Dan. Rev. Game B i o l . .7 ( 1 ) : 1-205. Sweeney, J . 1975. E l k movements and c a l v i n g snowcover. Ph.D. T h e s i s . C o l o r a d o S t a t e C o l l i n s . 95 p. S w i f t , R.W. Agric.  1957. The n u t r i t i v e e v a l u a t i o n Exp. S t n . B u l l . 615: 1-34.  as r e l a t e d to University, Fort  of  forages.  Pa.  180  T e l f e r , E.S. 1978. C e r v i d d i s t r i b u t i o n , browse, and snow in Alberta. J . W i l d l . Manage. 42: 352-361. Thomas, D.S. female British  cover  1970. The o v a r y , r e p r o d u c t i o n , and p r o d u c t i v i t y o f C o l u m b i a n b l a c k - t a i l e d d e e r . Ph. p. . T h e s i s . U n i v . o f C o l u m b i a . 211 p.  T i n b e r g e n , N., M. Impekoven, and D. F r a n k . 1967. An on s p a c i n g - o u t a s a d e f e n c e a g a i n s t predation. 28: 307-321.  experiment Behaviour  Traczyk, T. and H. T r a c z y k . , 1977. Structural characteristics o f h e r b l a y e r and i t s p r o d u c t i o n i n more important forest c o m m u n i t i e s of P o l a n d . E k o l . P o l . 25: 359-378. Turner, F.B., E . I . J e n n r i c h , and J.P. W e i n t r a u b . 1969. Home r a n g e s and body s i z e o f l i z a r d s . E c o l o g y 50: 1076-1081. Van B a l l e n b e r g h e , V. And J.M. Peek. 1971.. Radio-telemetry studies o f moose i n n o r t h e r n M i n n e s o t a . J . W i l d l . Manage. 35: 63-71. Van W i n k l e , W. 1975. Comparison o f s e v e r a l p r o b a b i l i s t i c r a n g e m o d e l s . J . W i l d l . Manage. 39: 118-123.  home  Verme, L . J . And J . J . Ozoga. 1971. I n f l u e n c e o f w i n t e r weather on w h i t e - t a i l e d d e e r i n upper M i c h i g a n . Pp. 16-28 In : A.O. Haugen ( e d . ) . Proc..Snow and I c e Symp. I n R e l a t i o n To W i l d l i f e And R e c r e a t i o n . 280 p. Verme, L . J . And D.E. U l l r e y . 1972. Feeding and nutrition of deer. Pp. 275-291. In : D.C. Church ( e d . ) . D i g e s t i v e p h y s i o l o g y and n u t r i t i o n of ruminants. Dep. Anim. S c i . Oregon S t a t e U n i v e r s i t y . V o l . 3. 351 p. Wallmo, O.C. And E.B. G i l l . 1971. Snow, w i n t e r d i s t r i b u t i o n , and p o p u l a t i o n d y n a m i c s o f mule d e e r i n t h e c e n t r a l Rocky Mountains. Pp. 1-15 I n : A.O. Haugen ( e d . ) . P r o c . Snow And I c e Symp. I n R e l a t i o n To W i l d l i f e And R e c r e a t i o n . 280 p. W r i g h t , E . and L.N. S w i f t . 1942. M i g r a t i o n c e n s u s of mule d e e r in t h e White River region of northwestern Colorado. J. W i l d l . Manage. 6: 162-164. Young, C. 6-12.  1977.  Cut along the dotted  line;  Forestalk  1  (4):  181  Appendix  I.  forested  plant  Actual  Estimates  of  associations  heights  were  shrub and  annual serai  stages  plant  associations  SC = Sword  F e r n - W e s t e r n Red  Hemlock,  WP = W e s t e r n  Fir  - Western  Western  Hemlock,  Hemlock,  available i n  during  summer.  used f o r s t e m s o v e r 30 cm. F o r stems  t h a n 30 cm t h e mean stem h e i g h t was s e t a t 0 Forested  growth  :  #  10,  and  VS = V a c c i n i u m - Skunk  Cedar,  DW = Deer  - Douglas-fir,  AT = A m a b i l i s  30  Cabbage,  AW  = Amabilis  SW = S a l a l  F i r - Twisted Serai  -  Stalk,  MC = M o u n t a i n Hemlock - C o p p e r b u s h .  Immature  N = Newly L o g g e d ,  S = Shrub, C = C o n i f e r .  H = Herb, F = F e r n ,  cm.  F e r n - Western  Hemlock - P l a g i o t h e c i u m , SD = S a l a l  less  Stages :  182  . . Association or S e r a i Stage  Annual Growth ( kg ha ) For V a r i o u s Mean H e i g h t s Of Stems < 30 cm  L  P l o t No.  0 cm  10 cm  30 cm  1005.7 700.7  1095.6  VS  7  974.2  VS  20  VS  41  666.3 136.8  SC  13  10.1  10.5  11.6  DW  19  16.3  18.0  25.2  WP  3  t  .1  WP  39  1.1  1.4  .3 2.8  AW  10  5.7  6.4  8.8  AW  18  4.6  5.3  8.0  AW  44  261.5  269.7  299.0  AW  45  154.2  159.8  182.4  SD  2  297.4  359.4  521.3  SD  5  575.3  643.9  823.3  sw  4  121.8  218.0  471.8  sw  6  190.4  218.1  302.0  sw  8  239.5  253.6  296.O  sw  17  19.6  21.4  28.6  sw  46  160.5  164.4  191.5  156.8  800.4 209.4  183  _1  I ^ A ^ , H A «  or S e r a i Stage  P l o t No.  Annual Growth ( kg ha ) For Various Mean Heights Of Stems < 30 cm 0 cm 10 cm 30 cm  AT  1  309.5  323.5  369.4  AT  28  992.1  1018.0  1104.8  AT  29  263.6  279.4  331.1  MC  32  364.4  406.5  545.4  MC  33  471.1  531.0  727.6  38  5.1  5.4  7.4  9  1.5  5.7  21.8  .3  1.6  .3  1.3  H H  11  H  14  t t  H  15  .1  .5  1.7  H  16  t  t  .2  H  21  13.3  20.1  52.9  H  23  67.3  83.8  137.9  H  24  11.3  12.1  17.2  H  30  7.2  55.5  214.0  H  31  2.7  3.3  5.8  12  184  Annual Growth ( kg ha ) For V a r i o u s Mean Heights Of Stems < 30 cm  Association or S e r a i Stage  P l o t No.  0 cm  10 cm  30 cm  S  27  122.8  234.0  524.6  S  34  32.8  39.6  62.2  S  36  .1  .2  .5  S  37  7.4  7.5  8.1  S  40  2.9  3.0  3.5  S  43  268.7  290.0  361.8  C  22  12.5  12.7  14.1  C  257.0  469.4  1027.2  c  25 26  306.2  461.2  866.3  c  35  195.7  515.8  135L7  c  42  139.8  141.3  146.7  t = trace  

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