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Aspects of the ecology of black and grizzly bears in coastal British Columbia Lloyd, Kevin Alexander 1979

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ASPECTS OF THE ECOLOGY  OF BLACK AND  BEARS IN COASTAL BRITISH  GRIZZLY  COLUMBIA  by  KEVIN ALEXANDER B.Sc.  (Hons.),  University  LLOYD  of British  A THESIS SUBMITTED IN PARTIAL REQUIREMENTS  Columbia,  FULFILLMENT OF THE  FOR THE DEGREE GF  MASTER  OF SCIENCE in  THE  FACULTY OF GRADUATE {DEPARTMENT  We a c c e p t to  this  the required  UNIVERSITY  STUDIES  OF FORESTRY)  thesis  1975  as  conferring  standard  OF BRITISH  COLUMBIA  S e p t e m b e r , 1979 1979  In p r e s e n t i n g  this  thesis  in p a r t i a l  fulfilment of  an advanced degree at the U n i v e r s i t y of B r i t i s h the L i b r a r y s h a l l I  f u r t h e r agree  for scholarly by h i s of  this  written  make i t f r e e l y a v a i l a b l e  t h a t permission  It  for financial  i s understood that gain s h a l l  of  The U n i v e r s i t y o f B r i t i s h  2075 Wesbrook Place Vancouver, Canada V6T 1W5  pate  flr^r  I agree  r e f e r e n c e and  f o r e x t e n s i v e copying o f  permission.  Department  Columbia,  this  3  Columbia  not  copying o r  for  that  study. thesis  purposes may be granted by the Head of my Department  representatives. thesis  for  the requirements  or  publication  be allowed without my  ii  ABSTRACT  This of  the  study  was  grizzly  americanus)  conducted  bear  in  habitat field  use  (Ursus  coastal  management p u r p o s e s . of  other  sign.  telemetry.  and  constrained  the  logistic  were l o c a t e d a t o t a l  a  of  occurred.  42  times.  Movement  Columbia  occurs,  but  behaviour  of b e a r s  on  reliable other  method was  bears  separate  the  t r e e s and  not  register  in  a  black  found  tracks.  salmon area  i t  consumed  21  Two from  and  monitored  sp.)  different  was  bears  of  vegetable insects.  Marking  were f o u n d  which  in a grizzly  matter The  r e c o g n i z a b l e foods.  No from  The  or j o i n e d ,  separate.  other four  space  British  of b l a c k bears.  are  Five  bears  close together  toe  was  described.  individual  Scats c o l l e c t e d  and  use  and with  movements  coastal  criteria  very  fifth  does..  scats  f o u r black  in  those  into  of the coast.  the ground  m i d l i n e of the  88 p e r c e n t  (Qncorhynchus  bear  during  were c u t  to c o l l e c t  grizzly  tracks  i s that the  below t h e  September a v e r a g e d  Trails  subseguently  to distinguish  t r a c k s are e i t h e r  bear  1977.  collected  of t r a v e l i s unknown.  on  t r a c k s of g r i z z l i e s  criteria  b e a r s was  watersheds  whereas t h e t o e s i n b l a c k b e a r reliable  and  feeding  36 t i m e s  the e x t e n t  using t h e i r  toes i n g r i z z l y  ecology  the  E x t e n s i v e o v e r l a p i n the  between  (Orsus  on  difficulties of  bear  and  and  o f b l a c k and  grizzlies total  black  research  were t r a p p e d and  study  by  and  ecology  Columbia f o r both  grizzly  i n 1976  a s p e c t s of the  systematically travelled  Bears The  British  black  a r e a and  aretos)  Information  s t u d i e s conducted  the study  to i d e n t i f y  A track  toes,  less does  whereas  from  May  through  and  12  percent  bears The  i n the bulk  of  study the  iii  diet  came  ladyfern  from  eight  (Athyrium  a u s t r i a e a) ,  of  these  filix-femina),  huckleberry  foods: spiny  (Vaccinium (Oglopanax  salmon.  The  early  herbs  f e r n s , and  and  and  salmon  the green fat,  were  bear  while crude remained  the  plant  n o t consume. protein soluble model  and  was  the  and and  For  the animal  As  the  crude  presented  gross  ratio  was  d e c r e a s e d , and moisture  b e r r i e s t o the c o a s t a l  devil's  club,  and  in  the l e v e l s  grouped  into  bear.  t y p e s and  future  were d i s c u s s e d .  implications  of  which i t d i d of  crude  of c r u d e f a t ,  The  A  simple of  vegetation  t h e bear  management o f b e a r s  Columbia  part  content increased.  this  British  the  levels  The  the  decreased, ash  t h e s e t y p e s were d e s c r i b e d . to  crude  total  the  grizzly  As  and  lower  matured,  Fruits  protein,  content  energy  sedge,  which d i s c u s s e s t h e e n e r g e t i c i m p o r t a n c e  s t u d y a r e a was  study  of  and  September.  consumed t h a n i n t h a t  berries  fibre  and  moisture  salmonberry,  the f i b r e : p r o t e i n  c a r b o h y d r a t e , and  salmon and in  increased  which  (Rubus  insects,  consisted  of crude  sp.)»  (Dryopteris  salmonberry  i n August  mature, t h e l e v e l s  constant..  huckleberry,  fern  horridum) ,  summer d i e t  extensively  carbohydrates,  fibre  (Carex  t h e c u r r e n t y e a r growth o f s h r u b s .  used  foods  soluble  and  wood  sp.) ,  gpectabilis) , d e v i l ' s club spring  sedge  foods i n  of the r e s u l t s  of  on t h e c o a s t o f  iv  TABLE OF  CONTENTS-  ABSTRACT  i i  List  Of  Tables  List  Of  Figures  viii ix  Acknowledgements CHAPTER  xii  1 - Introduction  CHAPTER 2 - The  1  Study Area  4  2.1.  Location  4  2.2.  Topography  4  2. 3.  V e g e t a t i o n And  2. 4.  History  Climate  4 7  CHAPTER 3 - Movements And 3.1.  Introduction  3. 2.  Methods  H a b i t a t Use  9 10  3.2.1.  Trapping  And  3.2.2.  Tagging  3.2.3.  Daybed R e c o r d s  And  Immobilizing Monitoring  And  Typing  Of  Bears  10  Bears  11 12  3.2.4.. Measurement Of  3. 3.  9  Bear Food  Availability  Habitat  12  Results 3.3.1.  15 Movements Of  Bears  3.3.1.1.  Grizzly  3.3.1.2.  Black  3.3.1.3.  Inter-valley  3.3.2.  3.3.3.  ,  15  Bears  15  Bears  I m p o r t a n c e Of Sources  -  Of  20 Movement V e g e t a t i o n Communities  Bear Food  R e s t i n g Cover  26 As 27 28  3.4.  Discussion  CHAPTER 4 - M a r k i n g  32  Behaviour  . ...  36  4. 1.  Introduction  . 36  4.2.  Methods  38  4. 3.  Results  40  4.3.1.  D e s c r i p t i o n Of  Bear Marking T r e e s  4.3.2.  S e l e c t i o n Of  Marking  4.3.3.  Frequency  Of  Use  Of  4.3.4.  Distribution  And  Abundance Of  Trees Trees  By By  40  B e a r s ...  43  B e a r s ....  43  Bear  Trees 4.3.5. 4.4.  46  Ground M a r k i n g  Behaviour  48  Discussion  CHAPTER 5 - The  Use  C e n s u s Of  49  Of  Tracks  I n The  Identification  Bears  -  And 53  5. 1.  Introduction  53  5.2..  Methods  55  5.3.  Results  5.4.  Discussion  65  Food H a b i t s  67  6.1.  Introduction  67  6.2.  Methods  69  CHAPTER 6 -  6.3.  • •-•  57  6.2.1.  Scat  Collection  69  6.2.2.  Scat  Analysis  70  6.2.3.  Bear Food C o l l e c t i o n  71  6.2.4.  Nutrient  71  Analyses  Results 6.3.1. 6.3.1.1.  72 S e a s o n a l Use Spring  And  Of  Food  E a r l y Summer  72 73  vi  6.3.1.2. 6.3.2.  Nutrient  78  C h a r a c t e r i s t i c s O f Bear Foods  O v e r v i e w Of R e s u l t s  82  6.3.2.2-  Nutrient  89  Food  Content  Of Bear F o o d s  Availability  93  Discussion 6.4.1.  93  C o m p a r i s o n Of Food H a b i t s  With  Other  Areas 6.4.2.  93  Implications  Of H i b e r n a t i o n  To  Feeding  Ecology  . 99  6.4.2.1.  Introduction  6.4.2.2.  The Model  6.4.2.3.  Implications  CHAPTER 7 - A d a p t a t i o n CHAPTER  -—  Of The Model  List  Of B e a r s To E n e r g y  Availability  I I : List  119 o f common  and l a t i n names used i n  III:  IV:  Appendix  V:  found  during  Seasonal  shifts  Ahnuhati  black  1976 Appendix  129  of species  watershed Appendix  i n the Ahnuhati  1976 and 1977 .  131  by t h e use o f f o o d s i n and g r i z z l y  bears  during  and 1977  Nutrient  values  . . . . 136 o f b e a r f o o d s by s e a s o n  Example o f d e t e r m i n a t i o n from  .... 108 113  thesis Appendix  99  104  8 - Management Recommendations  I:  ...  101  LITERATURE CITED Appendix  82  6.3.2.1.  6.3.3. 6.4.  Summer And F a l l  the energy a v a i l a b l e  salmon  .... 141  o f net energy i n coho 148  vii  Appendix  VI:  Body  measures and  particulars  f o r bears captured  Ahnuhati during Appendix  VII:  immobilization  1976  F r e q u e n c y and p e r c e n t understory  species  i n the  and 1977  149  c o v e r of  i n 14  plant  communities i n the Ahnuhati watershed, 1977  150  viii  LIST OF TABLES  Table  1: A v a i l a b i l i t y watershed,  Table  Ahnuhati  3: F r e q u e n c y species  food  in  the  Ahnuhati  1977  29  2: C h a r a c t e r i s t i c s the  Table  o f bear  of  trees  watershed  marked by b e a r s i n  ...........................  of occurrence o f among  18 b e a r  markinq  trees  by  tree  i n the Ahnuhati  watershed Table  44  4: C h a r a c t e r i s t i c s o f  grizzly  and  black  bear  paws Table  58  5: F r e q u e n c y  of  characteristics from Table  42  grizzly  6: The r a t i o  occurrence  of  which d i s t i n g u i s h  two  paw  black  bears  bears  of n a i l  distinguishing  61 length  t o forpad width  characteristic  as a  o f bear t r a c k s  64  ix  LIST OF Figure  1:  The  study  Figure  2:  The  upper A h n u h a t i  Figure  3:  Locations  Figure  4:  FIGURES  area  5 River  of a d u l t female g r i z z l y  No.  21  the  Ahnuhati watershed  and  Locations No.  7P  6  her  three  cubs during  bear 1977  16  of a d u l t female g r i z z l y  during  1977  in  i n the  bear  Ahnuhati  watershed ..................................... Figure  5:  Locations No.  Figure  6:  7C  and  7:  her  cub  Ahnuhati  watershed  Locations  of a d u l t  No.  during  12/22  Ahnuhati Figure  of a d u l t female g r i z z l y  Locations in  the  during  1977  bear  in  the  -  18  male g r i z z l y  1976  and  1977  bear in  watershed  the  ...  o f s i g h t i n g s o f unmarked  Ahnuhati watershed  during  19 bears  1976  and  1977 Figure  8:  21  Locations No.  20  9:  Locations No.  18  10:  subadult 1976  and  of  1976  in  the  and  female black 1977  in  bear  the  watershed of a d u l t  No.  during  watershed  1977  bear  22  subadult  Locations 19/23  female black  watershed  during  Ahnuhati Figure  of  during  Ahnuhati Figure  17  23 male b l a c k  1976  i n the  bear Ahnuhati 24  X  Figure  11:  L o c a t i o n s o f s u b a d u l t male b l a c k b e a r No.  4 d u r i n g 1977  i n the  Ahnuhati  watershed Figure  12:  25  V e g e t a t i o n communities  i n the  bottom  watershed  of the Ahnuhati  valley map ^vv  Figure  13:  Distribution salmon  and  Ahnuhati Figure  14:  Figure  15:  Ahnuhati  bear Figure  16:  watershed  Frequency  Figure  17:  scar  d u r i n g 1976 on an  Distribution  and  1977  ........  41 of s i z e c l a s s e s  t r e e s at the time of bear marking  of  of m a r k i n g  measured  47  o f bear t r a c k s  and  a sample d a t a  which  were  collection  sheet Figure  18:  Two  56  features  grizzly Figure  19:  F i g u r e 20:  45  t r e e s i n the  watershed  Parameters  30  Amabilis f i r i n  study area  distribution  marking  Ahnuhati  spawning  s u b a l p i n e s e d g e meadows i n t h e  A bear-caused the  o f c o n c e n t r a t i o n s of  pocket  which  distinguish  black  from  bear f o r e p a w s  Seasonal s h i f t s i n the black  and  study  a r e a , 1976  Seasonal  grizzly  62 use  bears i n the  and  grizzly  study  a r e a , 1977  by  Ahnuhati 74  s h i f t s i n t h e use  black  of food  of food  bears i n the  by  Ahnuhati 76  xi  Figure  21:  Seasonal  shifts  concentrations spectabilis) Figure  2 2:  Seasonal  wood f e r n Figure  23:  Seasonal  of salmonberry  (Rubus -  ..  shifts  concentrations lady fern  i n the n u t r i e n t  . .... i n the n u t r i e n t o f combined  (Athyrium  concentrations  samples o f  f i l i x - f emina) and  (Dryopteris shifts  83  spiny  austriaca)  84  i n the n u t r i e n t of d e v i l ' s club  (Q£lo£anax  h o r r i d urn) Figure  24:  Seasonal  85 shifts  concentrations (Lysichitum Figure  2 5:  Seasonal  of skunk  cabbage  americanum)  shifts  concentrations huckleberry  i n the n u t r i e n t  86  i n the n u t r i e n t o f combined  (Vaccinium  samples o f  o y a l i f o l i u m and  V. a l a s k a e n s e ) Figure  26:  Seasonal  shifts  concentrations Figure  27:  habits 28:  i n the of sedge  nutrient (Carex sp.)  Comparison o f c l a s s e s of bear Ahnuhati  Figure  87  88  foods i n the of  food  o f bears i n c o a s t a l ecosystems  Reduction into  with e x i s t i n g s t u d i e s  -  94  of n u t r i e n t s a v a i l a b l e i n food  the net energy  f o r f a t production  ........  105  xii  ACKNOWLEDGEMENTS  It  is  with  have a s s i s t e d supervisor. support the  pleasure that  with t h i s Dr.  Fred  study.  Chambers, D a r y l l Funding,  provided Hebert,  Fish  and  the  University  my  of  Service.  Funds  Hazelwood  of  the  lab  B.C.  like  to  gratefully  who  thank  my  British  David  analysis  and  Shackleton.  Forest  provided  and  David  the  Products,  the Canadian  were  Branch  Allan  were p r o v i d e d by  Columbia  Columbia,  Parks  acknowledge  c o m m i t t e e members: D r s .  transportation  British  for  I  Chuck J o n k e l , and  W i l d l i f e Branch,  many p e o p l e  f o r h i s g e n e r o u s a s s i s t a n c e and  study.  by  e q u i p m e n t , and  B.C.  I would f i r s t  Bunnell,  i n a l l phases of t h i s  assistance  I acknowledge the  Wildlife  by  Grant  Shackleton  of  U.B.C. I  a p p r e c i a t e the  Ellis,  Susan  B.C. to  field Dr.  the  and  Hebert,  Murray Shunter,  Smith,  Fish  assistance that  Fleck, Daryll  Leigh-Spencer, Bud  field  Graham  Wildlife  and  Turnbull, Branch  in to  Dick  Beames p a t i e n t l y animal  grateful  interpreting h e r I am  Ellis,  Barney and  Lay,  Sally  Smith. Larry  Johnson  of  provided valuable l o g i s t i c  directed  literature.  Hamer p r o v i d e d u s e f u l c r i t i c i s m am  K o s o v i c , Dan  Rick  the  support  crews.  domestic  I  Bob  I r e c e i v e d from  grateful.  Ms.  Daryll  Herrero  and  f o r h i s generous  David  assistance  the t h e s i s  t o thank F r e d B u n n e l l ,  Hebert  of  6.  C a t h i Lowe t y p e d  I would l i k e  Susan F l e c k , and  into r e l e v a n t areas  Steven  of Chapter  to Alton Harestad the data.  Dr.  me  f o r the  moral  support  and Rick and  xiii  faith  which they  Above a l l , joined  provided  I would l i k e  t o thank  i n a l l phases o f t h e study  observations. figure  long  months o f o u r l i f e  frequently  study.  Susan F l e c k  who  cheerfully  and c o n t r i b u t e d many  valuable  I am g r a t e f u l f o r t h e many h o u r s o f h e r l a b  and  mine.  throughout t h i s  drafting.  f a r from  She was a c o n g e n i a l companion i n the rainforest  ideal.  This  study  when  work  during the  conditions  i s a s much h e r s  were  as i t i s  1  CHAPTER  Through the ages, the place  symbol  of  carefully 1976;  world, a l l  the  grizzly  now  1976;  local,  with  provincial,  research recently for  (Herrero  potential  1970,  between and  and  ongoing  i t focused  is  us  to  impact  1977  and  occur, of  and  more  1978,  an  valuable  management  are 1971,  history  bears  has  People  level  of  been at  the  presently  only  o f human d e v e l o p m e n t  (see  of  a century  Bear  but  1978).  information types  into  (Cowan 1972).  than  Smith  then  a  The  1955).  species  special  Martinka  1977).  Tevis  impacts  various  and  international  for  on t h e  as  1976;  men  a  farther  persist  Servheen  (Storer  national,  been  example J o n k e l  management  occupied  takes  will  f o r the f u t u r e of t h i s  has has  bear  J o n k e l and  conflict  are concerned  has  1  as t e c h n o l o g y  the e c o l o g i c a l r e l a t i o n s h i p marked  bear  t h a t i s wild, i f i t s ecology  considered  Kurten  INTRODUCTION  grizzly  i n Man's mind and,  artificial  1 -  is  If  effective  needed  development  on  on  the bear  p o p u l a t i ons. General consistent aspects  ecological over  of  wide g e o g r a p h i c a l  their  ecology  Craighead  e t a l . 1974,  G l e n n and  Miller  i  Common and  p a t t e r n s o f b l a c k and  vary  Pearson  areas.  grizzly  However,  (compare B u n n e l l  1975,  Nagy and  names a r e  given  i n Appendix  I.  are  particular Tait  R u s s e l l 1978,  1977).  latin  and  bears  1978, and  2  In on  the  c o a s t a l ecosystems of  the b a s i c ecology  This  thesis  1976.  Field  October, The use,  work was  and  a  and  study  December o f  1976;  ecology.  The  behaviour together  the  bears  periods May  bears  emphasis  that  also  was  are  of Smith  May  bears  on  is  food  i n March  to  of  September, 1977.  habits, habitat  in coastal  British  h a b i t s and  feeding  field  discussed.  and  This  (1978), p r o v i d e  lacking.  t o October of  d e s c r i b e the food grizzly  information  initiated  s e p a r a t i o n o f b e a r s i g n i n the  of with  major  grizzly  March and  movements o f b l a c k and The  Columbia,  which was  done d u r i n g  o b j e c t i v e s were t o  Columbia.  for  presents  and  primary  of b l a c k  British  marking  information,  a preliminary  basis  management d e c i s i o n s . This  ecology  study but  primarily  the  they As  basic  e m p h a s i s from  because  Information B.C.,  examined a s p e c t s  was  collected  a r e an  the  appear on  integral  study  limitations  fluctuating  they  the  river  p a r t of  working  water  in  levels,  the  t o September  May  watershed o n l y .  The  expected coastal  to  vary  mobility.  results between  was  and  on  bears the  grizzly  grizzly to  ecology  in  habitats.  dense r a i n  forest,  results  (1976,  1977)  in  watersheds as c o n d i t i o n s change.  bears.  and  rugged  are l i m i t e d  thesis  years,  to  Rapidly  f o r the  this  s e a s o n s , between  coastal  i n response  coastal  presented  bears,  of g r i z z l y  The  bear  disturbance.  because,  i t s s c o p e narrowed  our  of  black  more s e n s i t i v e  topography r e s t r i c t e d period  outset  black  progressed, of  of both  and  to  Ahnuhati can  be  between  3  The  reader  objectives  i s introduced to s p e c i f i c  of t h i s  study  a t the beginning  aspects of e a c h  and  specific  chapter.  4  CHAPTER 2 - THE  2-1.  Location  The  s t u d y was  c o n c e n t r a t e d w i t h i n the  which f l o w s i n a s o u t h - e a s t e r l y British  Columbia  Inlet  is  Knight I n l e t  2.2.  112 88  km  km  Ahnuhati  rugged.  km  inland.  inland  direction 33«;  north The  into  latitude of  River  Knight  Inlet,  30»).  Knight  and  extends  126°  Vancouver  Ahnuhati  valley  River  empties  into  ( F i g . 1) .  narrow,  common  ( F i g . 2).  than  a  river  and  R i v e r watershed,  E l e v a t i o n s range  deep,  and  flat  The  kilometer thus  influence described  2.3.  the  320  Ahnuhati  Topography  The  The  ( l o n g i t u d e 50°  located  approximately  is  STUDY AREA  of by  vegetation  glaciation  on  drainage, encountered  and  hemlock  is  outcrops  are  i s realized  in  be  free  topography  is  climbed.  apparent  and  less the The is  climate w i t h i n the c o a s t a l western zones  associated elevation.  i n the  rocky  valley  (1974).  area l i e s  are  The  range,  distance perpendicular to  w a l l s cannot  V e g e t a t i o n and  mountain  large  relief  most v a l l e y  Russell  study  average  horizontal  many i n t h e c o a s t  50 t o 2386 m.  b o t t o m e d and  full,  of  from  like  study  of K r a j i n a with  The  area  (1965).  differences  plants,  are l i s t e d  hemlock  Variations in in  landforms,  mammals, b i r d s , in  and  Appendix  and II.  fish The  5  Figure  2:  The  upper  Ahnuhati  r i v e r  7  climate  i s humid  v a r i e s from altitudes varies  throughout  152 cm i n most l o w e l e v a t i o n  (Weiss  p e r s . comm.).  between  frost  free  months  (June t h r o u g h  period  between  one  Knight I n l e t  until  appears  Knight that  Inlet.  bear  bear-man  month  (July),  Smith  (1978).  by h u n t i n g .  were n o t k i l l e d  in  ( p e r s . comm. t o  and European  relationships  p e r s . comm. t o  i n t h e Campbell hunters.  While  Prior  large  the  with  probably  were k i l l e d  f o r sport.  sport  i t appears grizzlies  While  t o 1930,  hundred"  of  t h e 4 0 men i n t h e upper  in  observed increases  R i v e r a r e a , and i n of the I n l e t change  timber  an  increase i n  associated  fish  Refuse  at  were  shot.  canneries  these  attracted bears  to estimate the  reported k i l l i n g  o v e r a 10 y e a r p e r i o d Knight I n l e t  sites  and  ( L . Darke  Concurrently,  i t i s not possible  h i g h a s W. G i b b o n s  a  this  Specifically,  camps and f l o a t i n g  B. S m i t h ) .  no  bears i n  numbers  B. Smith)  Exploitation  occurred.  increased,  b e a r s , many o f which  kill,  to four  d u r i n g t h e 1940's and 1950's and a marked  logging  The  1960, p r e s e n t b e a r p o p u l a t i o n s i n  grizzlies  harvesting  shore-based  approaching  are available f o r g r i z z l y  i n homesteaders  bears increased  salmon  after  W. G i b b o n s  American  temperature  p e r s . comm.).  h u n t i n g i n c r e a s e d i n t h e 1930's due t o  logging,  guided  statistics  from  have been i n f l u e n c e d  that  mean  September).  comments a r e p a r a p h r a s e d kill  in  varies  These  probably  average  (Weiss  History  area  a r e a s t o 432 cm a t h i g h  -28°C  2.4.  quantitative  in  The  precipitation  7 and 9°C w i t h maximum t e m p e r a t u r e s  38°C and minimums a p p r o a c h i n g  it  the year and a n n u a l  and s t a t e d  area a t t h i s  that time  "over many also  8  killed Inlet years  large  numbers  may have s u s t a i n e d although  i t  is  of  grizzlies.  heavy  mortality  unlikely  Ahnuhati  were v u l n e r a b l e  the  head  of t h e I n l e t .  the  I n l e t has h o t b e e n l a r g e  The g r i z z l i e s  that  during bears  of Knight  the  last  resident  40  i n the  t o t h e same d e g r e e a s t h o s e r e s i d e n t i n Since  1960, t h e l e g a l k i l l (Hebert pers.  comm.).  of  bears  in  9  CHAPTER 3 - MOVEMENTS  3-1.  Introduction  While  the  several  movements  interior  Craighead  ecosystems  1976,  Nagy  movements i n c o a s t a l The  continuing i n  as  t o the  grizzly  bear  potential  the  grizzlies  survival  habitat  relative  coastal habitat  logging  the  and  1975,  1978), d a t a on  i t s associated  h a s been  coastal  viewed  habitat,  I t i s not possible, different  of the  and e v a l u a t e t h e information i s  r e q u i r e m e n t s and t h e e x t e n t  of  by many  ecotype  To u n d e r s t a n d  on g r i z z l y  of n a t u r a l  f o r example,  clearcut  k n o w i n g what p r o p o r t i o n  I present  and  requirements  migration for  requirements f o rreproduction Specific  Pearson  of  s i z e s on  their  home  objectives  preliminary of  marked  resting  and  information bears  on t h e  and  feeding.  their Habitat  and d e n n i n g a r e n o t t r e a t e d .  o f t h e movements and h a b i t a t  use  were: 1.  to  document  grizzly 2.  to  the  natural  movements  bears i n c o a s t a l B r i t i s h  describe  availability  to  represent.  chapter,  movements  documented f o r  1978, J o n k e l  Columbia  1974).  impact  without f i r s t  this  of  of  of logging  ranges t h e c l e a r c u t s In  Russell  British  movements o f t h i s s p e c i e s . assess  been  ecosystems a r e l i m i t e d .  (see R u s s e l l  needed on g r i z z l y  have  (see f o r example  and  coastal  impact  bears  expansion  activities a threat  of  AND HABITAT USE  coastal o f bear  habitat food  o f b l a c k and  Columbia in  terms  of  the  study  10  3.  to  describe  and  grizzly  3.2.  bears  Trapping  Aldrich  capture  five  b l a c k bear  and  foot  as d e s c r i b e d by  grizzly  and  88  success  was  snare  were  24  fired  Equipment  Co.,  estimated  prior  administered kg  at  body  Fifty-eight  trail  and  immobilized  from  a  capture  given  disposable  Cap-chur  plastic data  and  all  gun  M99  (Palmer  Georgia).  per  45  respectively. was  average  trap night injured  (Etorphine  with  subcutaneously fitted  or  mg  M 50-50 2  mg and  administered These  intramuscularily 20  was  35  Penicillin  trauma.  with  f o r each c a p t u r e a r e  or  a t a dose o f  were r o u t i n e l y and  were  or Sernylan  antagonist  M99.  and  weights  weight,  administered  of i n f e c t i o n  syringes  M99  using a  Chemical  Body  The  immobilized  one field  The  were n o t  either  kg body  1 mg  to  paw.  a dose o f  either  Immobilization  Bears  either  possibility  of  where one  t o i m m o b i l i z a t i o n and  weight,  grizzly  (Phencyclidine hydrochloride)  (Dexamethalsone 21-phosphate) the  One  sets  (1977)  trapping bears.  with  Douglasville,  45 kg t o r e v i v e b e a r s  to reduce  Flowers  percent  hour p e r i o d .  (Diprenorphine hydrochloride)  were  cubby and  bears.  on t h e s n a r e d  or Sernylan  5 cc d a r t  Azium  black  (1971) and  t r a p n i g h t s per  s e t f o r one  hydrochloride)  per  coastal  bears  i n both  four black  trails  beyond s m a l l l a c e r a t i o n s Bears  used  Cowan  were r e c a p t u r e d .  trap  per  by  resting  immobilizing  J o n k e l and  cutting  one  for  s n a r e was  h o u r s were s p e n t  is  habitat selected  Methods  3.2.1. The  the  gauge  drugs with  needles.  g i v e n i n Appendix  VI.  11  3-2.2.  T a g g i n g and  Bears mounted  were on  Materials, these the  fitted  wide  with  nylon  Inc.,  collars  monitoring bears  failed  attached  of  with  an  measurements measures  ear  and  occasions  installation. or b o l t e d  wide T i g e r - w e b  to  other  removed  the  base  i n p l a c e and n y l o n webbing  of  the  ear. VI.  a  15 was  Body Not a l l  were r e c o r d e d f o r a l l b e a r s .  the  air  Materials). antennas  with  Both  were  antenna pole  tag  three  Wildlife  Bears  weights a r e i n c l u d e d i n Appendix  Radio-instrumented and  were r i v e t e d 3 cm  62901).  on  after  radio-transmitters  LP-2280-MD,  Illinois  shortly  coloured,  powered  (Model  on t h r e e o c c a s i o n s and  transmitters  length  webbing  Carbondale,  Transmitter-collars cm  lithium  and  installed  upstream  a  null  used  (Cushcraft  b e a r s were t r a c k e d model and for  TRX-24  Model 23 mobile  Communications) a t base  both from  radio  the  receiver  ground  (Wildlife  Hy-Gain  three  element  Yagi  tracking.  An  11 e l e m e n t  Yagi  was  mounted on  a 7 m aluminum  camp t o d e t e r m i n e w h e t h e r a b e a r  was  o r downstream o f camp.  Monitoring consuming  from  because  the  ground  and  both  t h e b e a r s moved g r e a t e r  c a p a b l e o f moving and b e c a u s e radio-signals  was  gave  the  d i s t a n c e s than  numerous r o c k c l i f f s  unreliable  overcame t h e s e p r o b l e m s ,  but c o s t s  to l e s s  per month.  than t h r e e hours  difficult  results.  restricted  Aerial t h e time  and  time  I  was  reflected tracking available  12  B e a r s were l o c a t e d by t r a p p i n g , using  five  The  sighting,  or  triangulation  r a d i o - l o c a t i o n s o r more.  location  data  are  not  treated i n a detailed fashion  because o f t h e s m a l l  sample s i z e and b e c a u s e a t t e m p t s t o  radio-collared  were o f t e n  of  bears  were t h e r e f o r e  3.2.3.  The type, for  bears  Daybed  location,  unsuccessful  date, and  daybeds e n c o u n t e r e d  unknown.  associated distance during  s i g n , canopy c l o s u r e ,  from  the  plots  composition plot  with  height  which  of  each  a radius  f o r that  Height  had  o f 5.6  plot  common site  particular  be  center.  using The  from  o f 100 m . 2  and t h i s used  to  nested species  circular  The  average  height  was used t o  estimate  species:  Area  1.8  m  10  m  2  0.5 - 1.49 m  2.5 m  20  m  2  1.5 - 2.99  m  3.5  m  40  m  3.0 - 4.99  m  5.0  m  80  m  m  of  a  Radius  0.0 - 0.49  recorded  and t y p i n g  were s a m p l e d  was d e r i v e d  m and an a r e a  would  s i g n were  availability  areas  was r e c o r d e d  size  class  a  sampling  o f each s p e c i e s  determine which cover  and n o n - f o r e s t e d  feeding  habitat  study.  3.2.4. Measurement o f b e a r f o o d habitat  circular  and t h e w h e r e a b o u t s  records  substrate,  Understory  locate  2  2  percent  13  5.0 The  +  percent  estimate. slope,  m cover  At e a c h  aspect,  stoniness, cover, feeding,  recorded site,  any  crown  species.  If  a  only  food  bear  these  parts  c l a s s of the food  Bear  food  counts  ( e . g . no b e r r i e s frequency  were  vegetation  visual  location, regime, vegetative  ( i . e . evidence  of  p a r t of each  at a site,  The  size  of  food  then  the e d i b l e  the  area  from  was d e p e n d e n t on t h e h e i g h t  concerned. were c o l l e c t e d , c o u n t e d ,  were  not o b t a i n e d  when f o o d s  i n September p l o t s ) .  and  weighed.  were  senescent  Average p e r c e n t  were c a l c u l a t e d  was c o n f i n e d t o a r e a s  sampled,  b u t no r e p l i c a t e characterize  cover  f o r a l l species i n  of the v a l l e y  One  p l o t s were  vegetation  were g r o u p e d t o g e t h e r .  similarity  Dombois  activity  recorded  and T a n o c k t e u c h r i v e r s .  for  -  total  consumed a s p e c i f i c  Ahnuhati  understory  a  and each  type.  Sampling  To  was  foods  of occurrence  2  moisture  percent  plant occurred  plants  Samples o f bear  species  m  trail).  p a r t s o f t h e p l a n t were c o u n t e d . which  each  closure,  associated  the bears  for  100  micro-topography,  s c a t , b e d , t r e e , and  Usually  m  t h e f o l l o w i n g were r e c o r d e d  terrain,  percent  and  5.6  subalpine  bottoms o f t h e  side-valley  was  obtained. types,  plots  These g r o u p i n g s  u s i n g Motyka's s i m i l a r i t y  index  with  similar  were  (ISMO)  tested  (Mueller-  and E l l e n b e r g 1974): ISMO =  Where MW  i s t h e sum  species  common t o p l o t s A a n d B, MA  values from  plot  of  (2 MW/MA + MB) 100  the  smallest  A, and MB i s t h e sum  percent i s t h e sum of  cover  value  of percent  percent  cover  of  cover values  14  from  plot  B.  vegetation  to  communities  20 1 m u s i n g  inch  air  the  ISMO > 50  by  (types)  46  cm  were a s s i g n e d  t o the  same  were mapped a t a s c a l e o f  e n l a r g e m e n t s of  Supplementary  delineation  communities. grid  46  photos.  assisted  with  communities.  Vegetation cm  Plots  The  area  of  20  chain  i n f o r m a t i o n from borders  o f e a c h community  field  between was  to  2.5 the  notes  vegetation  determined  with  a  overlay. The  with  communities i n Table  integers.  a r e composed following  The of the  Those t y p e s  primarily  the  communities  1 and  o f the  prefix  6 and  with  Appendix the  types  (e.g.  M6-7  prefix  VII M"  11  indicated  i s a mix  are  a r e mixes by  the  kg/ha o f b e a r  food  was (x)  K = (.^  Hx  u s e d t o d e r i v e an  which  occurred  in  10,000  C/D i  estimate each  K Bi/A )(.2 i  A^  1  K2 A = percent =  B  =  cover  number o f in  the  berries)  D = area  o f bear  (x) i n  plot  p l o t s c o n t a i n i n g bear  food  habitat  number  C = wet  integers of  7).  f o l l o w i n g procedure  K  which  composed p r i m a r i l y  type:  Where:  labelled  of  food  type plant  counted  parts  i n each  weight  (kg)  o f one  (m )  o f one  plot  2  (x)  (e.g.  plot  plant  part  shoots,  (Hx),  habitat  15  3. 3. . R e s u l t s 3.3.1.  Movements o f  3.3.1.1. I  located  groups bear Adult  female  September. was  9 km  at  et No.  28  1977  The  female  grizzly  failed  31  No.  located  b e a r s and  3 to 6). 1977  21  eight  two  A d u l t female  and  was  not  family grizzly  located  again.  c a p t u r e d with her  times d u r i n g  d i s t a n c e between h e r  was 4) .  male  12/22  her  S i n c e h e r r a d i o was p r o b a b l y was  between l o c a t i o n s  and,  was  five  intensive  to locate  No.  three  August  location  7P  was  believed times  aerial  functioning in  on  and  points  June  A d u l t male No.  elsewhere  12/22  the  was  within  (see f o r example  t o be 7P's  10  mate.  Hensel Female  during July  through  September  search f o r t h i s  b e a r on  August  i n the Ahnuhati  not  captured  s i n c e t h e c a p t u r e d a t e was  described  located An  bear  sedge meadow s i t e .  at the t r a p s i t e  a l . 1969),  (Fig.  she  and  breeding season  7P  bear  greatest  a lowland  with her the  (Figs.  c a p t u r e d on J u l y grizzly  grizzly  ( F i g . 3).  Adult 1977  individual  o f 35 t i m e s  20 was  on J u l y  bears  three  a total  No.  cubs  Grizzly  bears  both  and  Tanockteuch  b e f o r e and  watershed.  watersheds.  after  Thus,  20  this  the  date,  distance  i n F i g u r e 4 probably underestimates her  actual  movement. Adult August  3  September  female 1977 of  and 1977  grizzly was  No.  located  ( F i g . 5) .  7C five  was  captured  times  with her  during  cub  August  on and  16  Figure.3:  L o c a t i o n s o f a d u l t f e m a l e g r i z z l y b e a r N o . 21 c u b s d u r i n g 1977 i n t h e A h n u h a t i w a t e r s h e d . C  capture s i t e  •  August  v  camp  A  September  and her  three  17  Figure  4:  Locations of a d u l t female g r i z z l y i n the Ahnuhati watershed.  C  capture  v  camp  site  ^  June  ©  July  ' • A  b e a r No.  August September  7P  during  1977  18  v  camp  A  September  Figure  6:  L o c a t i o n s o f a d u l t male g r i z z l y b e a r No. a n d 1977 i n t h e A h n u h a t i w a t e r s h e d .  C  capture  v  camp  site  June  A  ®  July  •  August  12/22  during  1976  20  Adult 1976  and  male again  successfully for  1976  trap  and  grizzly on  bear  August  by  5 1976.  12/22 This  r a d i o - t r a c k i n g , but  1977  was  captured  bear 13  was  on  June  24  never  located  l o c a t i o n s were  obtained  from s i g h t i n g s , h i s c h a r a c t e r i s t i c  tracks,  and  l o c a t i o n s ( F i g . 6). The  other  movements o f a l l o f t h e (Figs.  grizzlies  3  to  intensive  study  3.3.1.2.  I 11).  trapsite No.  20*3  She  left  and  of  den the  1976  the  bear  July  No.  at  of  18  ( F i g . 9).  She  Adult  male b l a c k  bear  and  again  in July  1976.  his  collar.  collar  area.  were known t o s h a r e  o f 41  (probably  the  times a  (Figs.  8 to  subadult)  was  13 o c c a s i o n s i n 1976  bear  accompanied  No.  our  approach.  The  l o c a t e d w i t h i n a 4 ha  occasions  his  20  l o c a t e d on  area  in  ( F i g . 10).  No.  and  captured  times  study  nine  area  on  20  March  and  at  the  area  of  19  1977.  shortly thereafter.  Female b l a c k  He  intensive  a total  second b l a c k  was den  were u s i n g t h e  bear  A  left  each  t r a p i n f o r m a t i o n showed t h a t  black bears  black  (Fig. 8).  overlapped  bears  four  in July  bears  ( F i g . 7) .  Black  Female  1977  The  grizzly  other adult g r i z z l i e s  area  located  captured  6).  ( c o u n t i n g cubs)  In a d d i t i o n , t h r e e  and  No.  of  was  (probably  1976  and  was  removed h e r No.  refitted  19/23  During with  a  subadult)  l o c a t e d on  nine  was  captured  this  period,  T h i s bear  was  located after  probably  malfunctioned.  1977.  i n June of 19/23 and  1976  removed  located  August o f  Subadult  was  subsequent  c o l l a r i n J u l y of  a second c o l l a r  not  also  black  11  1976 bear  21  Figure  7:  Locations watershed ©  grizzly  v  camp  of s i g h t i n g s o f unmarked b e a r s d u r i n g 1976 and 1977. bear  O  black  bear  i n the  Ahnuhati  22  Figure  8:  L o c a t i o n s o f s u b a d u l t f e m a l e b l a c k bear No. a n d 1977 i n t h e A h n u h a t i w a t e r s h e d . C  capture s i t e  s  May  •  camp  A  June  probable M a r c h  '  densite  ©  July  1 9 7 7  •  August  A  September  20 d u r i n g  1976  23  C  capture  v  camp  site  A  June  ©  July  •  August  Figure  10:  L o c a t i o n s of a d u l t male b l a c k bear No. i n the Ahnuhati watershed. C  capture  v  camp  site  A  June  ©  July  •  August  19/23  during  1976  25  26  No. 4 was in  the confined The  as  captured  several  Several  tracks  height  1977,  and  o f land  small adult  Knight  followed  each  other  to  10).  (Figs. 3  were s i g h t e d  since  t h e y were by Cowan  (Fig. 7), they entirely  black  (1938).  On  Further  the decrease  in  On A p r i l  5  km  the  Kakweiken  with a h e l i c o p t e r over t h e where t h e  animal  i n t e r - v a l l e y movementi n a hanging  was  On J u l y 3 valley  of  were f o l l o w e d  up t o t h e  were m e l t i n g  and d i f f i c u l t  to follow  were h e a d i n g t o w a r d s  of the Ahnuhati River..  Hills  Creek,  On A u g u s t 20  1977,  was n o t  indirect  f o r i n t e r - v a l l e y movement  evidence  bear  tracks  in  Ahnuhati this  at is  moving t o t h e A h n u h a t i R i v e r  that for  this  Sim  One  Sim R i v e r  large  in  River  increase  time.  the the  present  a  No. 7P a p p a r e n t l y  1978), and t h e c o r r e s p o n d i n g  of  in  26 1976, t h e  4, i t s t r a c k s  in grizzly  the  interpretation  for  The t r a c k s  female g r i z z l y  tracks  area.  b e a r s move between  b e a r were l o c a t e d  July  watershed east  (Smith  grizzly  b e a r was o b s e r v e d  observation,  August  Inlet  confirming  grizzly  of land.  watershed. .  bears  i n t o the Kwalate watershed  Ahnuhati.  at l a s t  grizzlies  suggest that  grizzly  thereby  an a d u l t  height  is  the  o f an a d u l t  sighted,  black  occasions  I n t e r - v a l l e y movement  in  watershed  the  bears overlapped  phase p r e d i c t e d  observations  watersheds  of  on s i x  11.  black  reliably  the c o l o u r  3.3.1.3.  but,  other  n o t be s e p a r a t e d  colour;  the  shown i n F i g u r e  as t h e l o c a t i o n s  Although  in  area  l o c a t i o n s of the four  well  could  i n J u l y o f 1976 and l o c a t e d  the  during  in grizzly possible  g r i z z l i e s are  salmon  run.  The  27  salmon river  run  in  limits  the  p e r s . comm.). movement At  Sim  River i s s m a l l ,  bears  While  to  the  foregoing  writing,  w i t h a c o l l a r i n August  1978  n e a r t h e mouth o f Sim  Female  No.  the  Leigh-Spencer, pers.  3.3.2.  Sign  communities  are  importance  of  them  the  proportion  second  found  of  in  protection.  bear  fitted was  e a s t o f the Ahnuhati. was  Ahnuhati  located in  (Langin  and  communities as s o u r c e s of  a l l of  15  plant  the  Appendix  VII.  ways:  (i)  ( i i ) by r a n k i n g t h e  reveals  first  i n that  from  relative  the  by  can  ranking  be the  communities  to  in  type i n the study  method g i v e s  an i d e a  forage  bear food,  perspective  which  in.  contribution  supply of a v a i l a b l e  meaningful  relative  abundance o f b e a r f o o d s i n  f o r a bear  the  The  These  as s o u r c e s o f bear food  and  The  communities  Ahnuhati.  relative  1).  communities t o the t o t a l more  male g r i z z l y  to  i s most " p r o f i t a b l e "  probably  known.  the  in  t h e s e communities  according  method  not  h e r t h r e e cubs)  bottom  described  (Table  inter-valley  movement i s  11 km  t o t h e t o t a l amount o f f o o d  (kg)  community  west  valley  (kg b e a r f o o d / h a ) ,  area  Creek,  on a s e a s o n a l b a s i s i n two  communities  (Smith  comm.)..  o f b e a r use was in  assessed  an a d u l t  Importance of vegetation bear food  recognized  tributaries  suggests that  of t h i s  (and, p r e s u m a b l y ,  K w a l a t e w a t e r s h e d , 4 km  w a t e r i n t h e main  n e a r b a s e camp i n t h e A h n u h a t i  located  21  silty  clear-flowing  does o c c u r , t h e e x t e n t  the time of f i n a l  and  of  .The  of  the  and  is  habitat  28  The  absolute  habitat for 1,  type  (Table  a bear to forage M5-6-7, M6,  the  Ahnuhati  and  descending  most  the  7,  communities  in  Figure  and  1,  M5-7  by  9,  the  closure.  areas  bear  Sixteen  bear  ( F i g . 12,  1,  food map  F i g . 12).  largest  of  order:  communities  in  which  are,  in  pocket).  f o r a g i n g on b e r r i e s ,  The  the  four  descending four  plant  amount o f b e r r i e s  are,  M7.  o f s p a w n i n g salmon a r e  were  fished  h e a v i l y by  and  subalpine  bears  associated sedge  shown  and  as  tracks grass  13.  daybeds  beds  26  were  had  greater  area,  surrounding Frame the  examined  beds a v e r a g e d  than  Creek l e f t  communities  to forage i n are, i n  beds l o c a t e d d u r i n g  from heavy r a i n s .  15  by  cover  and  Olson  6  M5-6-7, and  crown c l o s u r e or h i g h e r the  and are  distribution  Resting  shown  f o u r communities  a b u n d a n t salmon c a r c a s s e s  The  All  The spring  concentration  Canopy c l o s u r e o f  100).  of the  of  (Table  M5,  These  Thirty-seven River.  12.  types  i s shown a l s o i n F i g u r e  3.3.3.  to  and  of high  13.  daybeds.  areas  biomass  habitat  are  s p r i n g are, i n descending  that bears  M7,  rankings  distribution  M5-6-7, 9,  order:  areas  evidenced  The  two  f o u r most p r o f i t a b l e  which c o n t r i b u t e the  descending The  The  i n during  6.  time  profitable  o r d e r : M6-7,  these  1).  largest  order:  During  of  i s shown i n F i g u r e  c o n t r i b u t e the  in  values  the  than  80  where t h e y  that bears that  (range  percent  b e d d i n g m a t e r i a l was suggesting  Ahnuhati  percent  r a i n y w e a t h e r had  (1974) o b s e r v e d  areas  59.4  i n the  black  were f e e d i n g  on  80  0  canopy percent  much  drier  seek  refuge  bears  at  salmon  and  Table 1.  A v a i l a b i l i t y of bear food i n the Ahnuhati watershed, 1977. A l l values are kilograms per hectare unless noted otherwise. Plant community type number  Food species  2 shoots of salmonberry  M7  M5-7  M6-7  H5-6-7  33.0ha  3.9ha  36.0ha  12.6ha  3.8ha  50. 3ha  Total (kg)  459.0  6.4  46.9  147.0  121.0  90.4  84.4  36,605  (15.6)  2.6  96.0  360.0  210.0  15.6  286.0  (223.0)  30,829  (16.6)  54.3  6.7  15.0  (163.0)  126.0  (166.0)  59.3  2  3  4  5  6  7  8  9  H5  2.14ha  1.29ha  U.19ha  26.8ha  76.5ha  20.0ha  32.5ha  0.81ha  30.0ha  tr  tr  tr  87.0  286.0  (16.0)  15.6  786.0  13.0  2.2  224.0  leaves of d e v i l ' s club sedge  m  1  1  8900.0  19,046  berries of salmonberry skunk cabbage  tr 62.4  1131.0  spiny wood fern  69.0  lady fern  3.4  20.3  tr  berries of d e v i l ' s club b e r r i e s of Alaska huckleberry leaves of Alaska huckleberry  tr  tr  berries of t a l l blue huckleberry leaves of t a l l blue huckleberry  tr  berries o f s t i n k currant  Total (kg) 3  Rank Sample s i z e (N)  21.8  3.2  21.0  24.8  5.0  101.0  3.1  45.8  5.9  53.0  *  (35.6) 192.0  18.0  0.6  14.0  tr  tr  tr  10.2  17,105  20,520  13  15  7  5  1  4  14  4  3  1  1  5  3  8  est.  *  SN  48.0  54.0  (41.9) (4.8)  0.5  .3  (kg o f food x area o f type)  18.8  7.8  (12.1)  not separated from Alaska huckleberry  18,629  237  (75.0)  (5.4)  7,927  SN  (54.3)  5,489  2.4  55.7  (50.3)  5,155  tr  5.6  tr  7.0  4,730  tr  6.4  tr  6.5  4,620  tr  0.4  tr  tr  2.0  1,491  tr  tr  tr  tr  1.6  1.152  7.0  tr  tr  2.1  536  19,248  4,412  2  10  6  est.  •  97.9  tr  SN = plants senescent.  2 » s c i e n t i f i c names given i n Appendix I  (73.0)  6.3  (14.4)  1.485  (67.4)  0.7  tr  tr  10.5  118.0  (2.4)  19.137  t r = trace 1 =  18.0  30.0.  12,808  13,803 13,438  442.0  tr  1,908 11 3  4.724  1,642  8,194  6  9  12  8  est.  1  est.  14,796  e s t . = estimates  3 = ( kg of food) x area o f type  est.  1  Figure  13:  D i s t r i b u t i o n of c o n c e n t r a t i o n s of spawning salmon s u b a l p i n e s e d g e meadows i n t h e A h n u h a t i w a t e r s h e d 1976 a n d 1977. Spawning  *  Camp  salmon  Subalpine  sedge  and during  meadow  31  sought the The in  cover of  daybeds  several  eight  u s e d by  respects  daybeds  in  was  apparently  situated m of  the  beds was  in  seven  cases,  of  scats  were n e v e r  60  cm  deep.  of bear  feeding  to the  more  than  the  uphill  side  formed  scraped  by  the  to  soil. of  The  was  12  not  observed  were n e v e r  scats  The  were s i t u a t e d i n 150  of  m  from  large  tree  and  side  and  of  the  associated  the  conifers,  roots.  ferns  one  differed  salmon r u n .  A l l eight  Bedding mosses. the  bear  daybeds  with  during  30  m.  the  them,  and  salmon  cases,  bedding  with  i n 10  these  cases.  beds.  the  substrate  n e e d l e s of c o n i f e r s and  recorded with  The  run  deciduous f o r e s t  understory associated  soil  Accumulations  Beds m o n i t o r e d  on  re-used.  measured a b o u t  greatest was  the  were  mature c o n i f e r o u s  sand i n  and  s a l m o n run  salmon run  t h i c k e t s of s a l m o n b e r r y .  typically The  used d u r i n g  daybeds l o c a t e d  river.  these  periodic basis  29  the  of  t o the  rains.  use.  both  usually  heavy  o f c o n i f e r s or  exposed  in  b e d s was  Daybeds  was  the  repeated  150  area  accumulations  In c o n t r a s t ,  within  flat  often  l a y on  large  suggesting  a  prior  e i t h e r needles  material  were  those  f o r e s t and  relatively  substrate  had  bears p r i o r  These d a y b e d s were on  the  This  from  located  mature c o n i f e r o u s river.  mature f o r e s t d u r i n g  distance  1 m long  observed from  by  1.3  m wide  daybeds t o  by  sign  32  3. 4.  Discussion  With bears  t h e e x c e p t i o n o f movements t o s u b a l p i n e sedge in  the  Ahnuhati  were  confined  E x t e n s i v e o v e r l a p i n t h e use of space 13  known g r i z z l i e s  (including  u s i n g t h e same v a l l e y Although  bottom  overlap  cubs)  to  meadows,  valley  bottoms.  o c c u r r e d among b e a r s ,  and f i v e  known  black  with bears  area.  occurs  in  o t h e r a r e a s , no o v e r l a p among  b e a r s as g r e a t a s t h a t o f t h e A h n u h a t i  h a s been o b s e r v e d ,  other  than a t c o n c e n t r a t e d food sources  a s salmon  River  (Stonorov 1972), Nagy  and  Stokes  1972) ,  o r s e d g e meadows and  Russell  suggested  (1978)  that  elk  on K o d i a k  m a l e s whose home r a n g e s data  several  among  five  male  in  the  black  The g r e a t e s t o v e r l a p t h a t and  (Atwell  f o u r female  overlapped  adult  shared a very s m a l l p r o p o r t i o n of t h e i r the  overlap  distribution isolated  observed  movements  data to o f f e r  of g r i z z l y  C i r c u m s t a n t i a l evidence  cited  movement o c c u r s i n c o a s t a l Ahnuhati,  b e a r s from  female home  and two  hills.  Their  using the  observed grizzlies  ranges..  was which  Perhaps  i s due i n p a r t  to the  to  the  topographically  involve  long  movements a l o n g  ( F i g . 13).  t h e home r a n g e  the  Swan  (Cole  a l . 1977).  grizzlies  (1975)  Ahnuhati  s e d g e meadows n e c e s s a r i l y  I have s u f f i c i e n t  in  the  of food, since  v a l l e y bottoms  of  in  et  b e a r s were a l s o  Pearson  two  a t McNeil  c a l v e s i n Yellowstone  Island  found  area.  adult  such  nearby  l o c a t i o n s observed  and  bears  in coastal  earlier  like  British  suggests t h a t  B.C. and t h a t  rivers  valleys.  only a preliminary  salmon  estimate Columbia.  inter-valley  spawning  i t , may be used  by g r i z z l y  The g r e a t e s t d i s t a n c e between  i n t h e Ahnuhati  areas  was 9 km f o r an a d u l t  radiofemale  33  grizzly  bear  bottom  (Fig- 3).  and  thus  rectangular. length  of  published  data,  ranges  (Pearson  1978,  and  has  a  1.5  and N o l a n  o f an a d u l t  male's  of  a  grizzly  bear  of a d j a c e n t portion  of  to  and Beecham  selection  the  (1977)  of female  home  and  Russell  home  that  and  (1976)  t h e movements and primarily  Hensel  i n salmon  (1972) spawning  on t h e m i g r a t o r y  habits  t o be i m p o r t a n t f o r b e a r s  over extremely t o spend  (Carex m a c r o c h a e t a ) Island.  rugged  They  stages  an a l m o s t  that  of  exclusive  as the primary  found  terrain  c o n s i d e r a b l e time  late successional  A t w e l l e t a l . (1977) f o u n d  brown b e a r s on K o d i a k  abundance,  and  variations  appear  s i t e s are possibly  f o r sedge  of bears i n the  areas.  r e a c h t h e s e a r e a s and they  subalpine lakes.  Thus,  probably  are governed  Berns  t o have a marked i n f l u e n c e  These  that  range,  Amstrup and Beecham  suggest that  s u b a l p i n e sedge meadows a p p e a r  preference  movements  o f b l a c k b e a r s i n Idaho  bears i n c o a s t a l  them.  from  Ahnuhati  distribution,  d u r i n g summer, a s t h e b e a r s t r a v e l  in  computed  rectangular  the  (1972) n o t e d  Kistchinski  the  watersheds.  and  to  as  the  or  i s the width o f  1976, Nagy  in  o f key f o o d p l a n t s .  The  as  T h u s , one would p r e d i c t  by t h e p h e n o l o g y  o f brown  considered  whose w i d t h  o f t h e major f o o d s p e c i e s .  times appear  be  elliptical  t o 2.1 t i m e s t h e a r e a  et a l . 1978).  may be r e l a t e d  Reynolds  habitat  could  likely  w i d t h o f 0.5 km, would be 13.5 t o 18.0 km.  significant  phenology  are  M a l e home r a n g e s ,  1975, P e a r s o n  portions  Ahnuhati  and  are  Russell  t h e home r a n g e  A  movement  (0.5 km).  minimum l e n g t h  includes  ranges  a r e c t a n g u l a r home r a n g e floor  which  home  The 9 km  the v a l l e y  the  Topography c o n f i n e s b e a r s t o t h e v a l l e y -  brown  food of bears  reached  densities  in  subalpine  meadows  o f 2.6 b e a r s p e r km  2  d u r i n g a 5 t o 6 week p e r i o d . The  relative  sources While  of  importance  bear  the actual  spring  food  has  importance  i s not well  development  of the various plant  could  potential  of plant  be r e d u c e d  because  of  sedge their  o f any p r o p o s e d  communities  to  by c o n s i d e r i n g t h e i r  bears  are  not  topographical isolation.  mining  on  relative  vulnerable  these  sites  in  of future value  t h e most v a l u a b l e c o m m u n i t i e s  meadows  as  management i m p l i c a t i o n s .  u n d e r s t o o d , t h e d e t r i m e n t a l impact  as f o o d s o u r c e s and p r e s e r v i n g Subalpine  communities  to  .  logging  However, t h e i m p a c t  should  be  considered  carefully. Circumstantial spawning  valleys.  the Kwalate,  the  from  Known g r i z z l y  Leigh-Spencer  Ahnuhati  by  may be used bear  suggests that by g r i z z l y  populations and F r a n k l i n  and  Hebert  (Hebert  unpubl.  therefore affect  data).  occur  to  Disturbance  and i n c r e a s i n g  A h n u h a t i , and a r e a s  consider  potential carefully  i m p a c t s on t h e s a l m o n controlled,  and  resource.  active  t h e salmon r u n u n l e s s i t i s l o c a t e d salmon s p a w n i n g  move  of the of  the  m o r t a l i t y but  Thus, development  i t , should  in  p e r s . comm.).  a l s o the bears o f adjacent watersheds. like  from  watersheds  not only the g r i z z l i e s  lowering productivity  salmon  bears  t h e s e a r e a s have t h e p h y s i c a l c a p a b i l i t y  salmon r u n c o u l d Ahnuhati  earlier  K a k w e i k e n , Sim, K l i n a k l i n i ,  (pers. o b s . and  to  cited  areas i n the Ahnuhati  surrounding  Grizzlies  evidence  i nthe  carefully  the  Development s h o u l d be  development  not occur during  more  2  a r e a s shown i n F i g u r e 13.  than  km  from  the  35  It during  appears the  important  that  period  bears  prefer  p r i o r to the  function  of t h e s e  mature t i m b e r  salmon r u n . sites  f o r daybed  Probably  i s s h e l t e r from  The  e x t e n t t o which s e r a i s t a g e s c o u l d  not  known.  serve  this  sites  the  heavy  most rains.  function  is  36  CHAPTER 4 - MARKING  4.1. The their  Introduction behaviour  habitat  of p h y s i c a l  is  and c h e m i c a l m a r k i n g  widespread  among mammals.  known e x a m p l e s a r e among mammals which  produce  squirrels, al.  Carnivora  scent d u r i n g marking  Eisenberg exhibit  behaviour  (1966)  marking  bears i s t h a t Like  transmit  a  the  lion  (Rudnai  generalized  handled  Meyer-Holzapfel  bears.  first  of trees  "odour"  when t h e b e a r r e l e a s e d observed  by  Hediger  Lindemann  mark  for  the  bears  wallowed i n u r i n e - d r e n c h e d  backs  against  behaviour  In the Ursidae,  trees.  Many  (Muller-Schwarze  1900's  behaviour of  their  stated  in  the  this  that  body.  that  of  1974) .  and  then  they The  who h a v e  mechanical  by an o d i f e r o u s a  Basle  type  (Davis  of  tree  (as  Zoological odiferous  i n t h e C a r p a t h i a n Mountains. dirt  et  marking  early  i ti s possible  was a c c o m p a n i e d  (1954) d e s c r i b e d bear  the  urine while rubbing  Gardens) .  brown  Burkhardt  w e l l known t o p e o p l e  [1946]  ground  many, i f n o t a l l o f t h e  from  (1957)  by b e a r s o f t e n  marten,  i n t h e marking  1973),  is  glands  e x t e r n a l scent glands  scent  bear  marking  that  behaviour.  apparently lack  obvious  1970; b a d g e r s ,  stated  characteristic  marking  possess  (e.g. beaver,  The major d i f f e r e n c e they  of objects i n  Most o f t h e w e l l -  h a s been r e c o g n i z e d s i n c e a t l e a s t  1919).  1964).  that  and mongooses, B o u r l i e r e  1967).  (Mills  BEHAVIOUR  rubbed  the cervids a l s o e x h i b i t  These their this  37  Wright served  (1909) m a i n t a i n e d  a  territorial  (Holzworth that  1930, K r o t t  marking  when t h e y respect could  that  t h e marking  function, and K r o t t  trees indicated  were l o c a t e d  at  not reach higher.  this  1963).  greater  bear  has  Mills  the l i m i t s  a  f o r the resident  but  o f t r e e s by  bears  been  denied  (1919)  doubted  of a d i s t r i c t  height,  they  or that, commanded  a n d s c a r e d o f f any i n t r u d e r  He r e g a r d e d  who  marking t r e e s as " p l a c e s o f  orientation". Murie such  (1954) s u g g e s t e d  as  place  Cahalane  (1947)  with  that  believed  irritation  Ground  marking  has  grizzly  been  bears  Holzworth result  which  consistently  that  of  I  by s h e d d i n g  stretching. with  insects  the winter coat  and  and  in  a series  o f depressions of  association  with  marking  documented a s one o f t h e m a r k i n g b e h a v i o u r s o f p e r s . comm.,  type  S u c h marks have been  along  by a b e a r  "pacing"  comm.,  regarded  as t h e  in a  visit  ritualistic  pers.  s t e p i n t h e same p l a c e s , e v e n t u a l l y r e s u l t i n g depressions  of  Jonkel  bears  a trail.  The A h n u h a t i  i ssufficient  t o create  d a t a show a  series  marks.  collected  ground marking  information  on  but  both  t h e m a r k i n g o f t r e e s and  d u r i n g t h e 1976 and 1977 f i e l d  do n o t p r o v i d e a l a r g e b a s e f r o m derived,  were m e r e l y a  and  were p l a g u e d  signposts,  whereby  a single  ground  be  they  rubbing  produces  trails  (Martinka  some  or that  that bears  caused  1930, Day 1957).  of  series  t r e e s might  stimulated rubbing.  s t e p s along bear  trees  of  f o r comfortable  the i r r i t a t i o n s  this  bear  such  scent posts o f the Canidae,  convenient  and  that  they  do  provide  which  new  further  seasons. hypotheses  description  The d a t a can of  be  this  38  behaviour. rest  Final  i n t e r p r e t a t i o n of the function  on c o n t r o l l e d e x p e r i m e n t a t i o n  presented  are o f f e r e d as a background  Specific 1.  2.  objectives  to  describe  for  marking  to describe by  3.  (Eisenberg  bears  to  of marking  1966).  f o r such  The  data  study.  of the study o f marking behaviour  t h e manner  will  were:  i n which b e a r s use h a b i t a t  the c h a r a c t e r i s t i c s of  trees  selected  of t r e e s  marked by  f o r marking  document  the  distribution  bears.  4.2.  Methods  During area  in  initial 1976,  reconnaissance  I  noted  the  study  t h e l o c a t i o n o f 43 b e a r marked  trees.  During subsequent systematic any  new  occurrences  g r o u n d marks other  t r a v e l over the  of v i s i b l e  marking.  were measured, and t h e i r  bear  substrates  and e x p l o r a t i o n  sign  of  area,  I  recorded  The s i z e and number  l o c a t i o n and p r o x i m i t y  recorded.  To  record  were e s t a b l i s h e d a t  the  bases  bear of  to  visits,  several  of  sand  marking  trees. The of  low f r e q u e n c y  marked t r e e s  frequently  suggested  than  of f i s h i n g  several  trees.  the  stretched  with  pitch  dislodged  that  suggested  Sections  line  o f f r e s h p h y s i c a l marks and t h e abundance  line  One  from  the l i n e .  bears  by were  were  the  across  the scar,  the tree.  physical  fastened  end o f the l i n e  marking  evidence  across  was t i e d  the  scar  into  rubbed a g a i n s t  more alone.  to a short  and t h e n p a s t e d  Bears that  trees  on  nail, place  the tree  39  Several The  age  rings  c h a r a c t e r i s t i c s were measured  of the  on  cores  tree  was  extracted  corrected  f o r the  t r u e age.  I estimated  by  the  rings  the  determined  the  s i z e of  by  recording  the  length  1.3  m  the  counting  above  of  marked  the  ground.  Ages  between t h i s  the  scar  on  estimated  tree  at the  time  second c o r e .  dimensions of  growth  were  height  the  age  trees.  annual  not  and  the  marking  and  the  at a comparable h e i g h t  of t h e  (dbh) , t h e  by  i n age  age  taken  I  at  m  between t h e  a second core  diameter  1.3  difference  difference  on  estimated  f o r 24  on  tree  number the  scar.  t h a t i t was  marked  I a l s o recorded the  scar,  of  the  and  its  height. To for I  evaluate  null  marking i n p r o p o r t i o n  recorded  the  species  Species  of t r e e s  itself  were  along  the  center.  sites  The  It  was  b e a r s use  forest  species.  their 20  I  2)  to the  not  trees  the 18  marked to  of  each  of  10  tree  species  the  forest,  18  sites.  marking  tree  t r e e s i n each d i r e c t i o n 4  m  from  marked t r e e s species  the  trail  i n Table  availability  3  was  of  18  trees.  evaluate  sizes i n proportion  b e c a u s e of t h e  within  l a r g e r than the  mean t r e e  possible  for  more t h a n  frequency  b e a r s use  occurrence  selected  which were n o t  applying  tree  to  s i m i l a r s i z e or  expected  by  (Table  of  hypothesis that  of  recorded.  trail  calculated  the  the  to  the their  differential  null  hypothesis  occurrence  that  within  growth r a t e s among  tree  40  4.3.  Results  4.3.1.  Description  Figure visited first the  14  by  Fig.  in  14) .  t h o s e of inner  about  1977. of  tree  cambium  scar.  Bears that  diameter Schaffer  the  an  the  the  hair l e f t  was  a  of  other  trunks  (66  trees  Montana,  w i t h few  (foreground  of  were s i m i l a r  to  the  all  branches  cm  (Table of  the  then  the  trail.  the  bear  licked  the  frequently  leave  pitch.  The  total  Some  trees  Bear  tree  scar.  dbh),  about t w i c e  2).  As  reported  marking  protruding  the  that  scar,  measurable  l a r g e s t t r e e examined  was over  i n a l l t r e e s examined.  show  marking  in  tree  grown back  mark f a c e d  trees  imbedded  the  in  the  This  was  been removed,  indicated  marking  small  d i d not  trail  the  marked t r e e  underhair  not  b a r k had  e x p o s e d , and  against  study.  had  the  f i r , which  w h i c h I examined  5 i n that  still  Amabilis  cambium  faces  scars  trees  trees  below  1.5  the by had  m  above  79  years  ground. The  mean age  (range 48 tree  No.  rub  and  (1971)  straight  5,  i t s muddy p e l a g e a g a i n s t  were c l a w e d , but 5 was  the  scar  a freshly  rubbed  No.  and  the  was  first  guard  1950 The  All bear  amount o f  No.  bears s e v e r a l times during  O b s e r v a t i o n s of  both  bear marking  shows b e a r t r e e  scarred  scar  of  to  117  of  years)  s c a r r i n g was  the  l a r g e s t scar  and  the  trees  20  on  mean h e i g h t  ( r a n g e 60  to  238  cm)  used  (Table  years  the  by  (Table  The  mean age  (range 5 t o  trees  of the  2).  b e a r s f o r m a r k i n g was  was  highest 2).  970  cm  38 2  marks on  of  years). (range the  the  first  Mean s i z e 10-3850  trees  was  of cm ) 2  139  cm  41  Table 2.  C h a r a c t e r i s t i c s o f t r e e s marked by bears 1n the Ahnuhati watershed.  i ree reference  DBH Scar age  Tree age  A t time o f marking (cm)  number  (years)  (years)  1 2 3 4 5 6 7 8 9 10 13 14 15 16 17 IS 19 20 21 22 23 24  14  48  29 29 24 38 13 25 17 5 21 20 N/A 21 N/A • N/A N/A  X . Range  _ _ 66 59 80 114 86 92 60 65 100 86 100 117 54 49 80 94  15 5  94 64.  19.7 5-38  79.4 48-117  A = Amabills f i r DBH = diameter a t b r e a s t  DBH (1977) (cm)  30.0  • 44.0 22.6 18.0 21.0 7.0  32.4 45.0 66.2 30.6 21.0 34.4 9.0  6.5 23.0 N/A 24.0 N/A ' N/A N/A  52.5 38.0 31.0 62.5 16.0 25.0  N/A 38.0  36.0 43.0  22.6 6.5-44  36 9-66.2  Tree s p e c i e s  A A A A A H A A A  A A A A A A A A A A A A  N/A N/A  P r o p o r t i o n o f t r e e s i n t h e Largest s c a r area t h a t were the same dimensions Area s p e c i e s as t h e marked t r e e 2 (cm) (cm )  -20 70 35 60 55 55 40 40 2' 5 45 20 20 75 20 20 70  — 55 30  • 36x90 9x25 60x10 60x30  3150 225 . 600 180  2x5 . 10x3 no scar 50x8 no scar no scar no scar 38x7.5 no scar 110x35  • 10 30 N/A 400 N/A N/A N/A 285 N/A 3850  970 10-3850  Highest point of  scarring •  (cm)  — — — . -— — — — 140 140  —60 — 200 120 N/A N/A N/A N/A N/A N/A N/A N/A  139 60-238  N/A » n o t a p p l i c a b l e height  ro  43  4-3.2-  Selection  Sample  sizes  in  not r e c o r d a l l types in  Table  fir  2.  o f marking  the following  3).  Of  amabilis western  fir,  r e d cedar.  significantly marking  of  therefore  western  reject  s t u d y a r e a were  the n u l l  occurrence within that for  amabilis  bears  occurrence  spruce,  Table and  f o r marking  of  The a v a i l a b l e  3)  amabilis  sitka  this  available. marking  trees  and  by  included  occasionally  o f a m a b i l i s f i r was  spruce  combined.  the tree  by b e a r s  and a c c e p t  when  than t h e frequency of  hypothesis that  the f o r e s t ,  were  11 were c h o s e n  o f marking  f i r are selected  I  s p e c i e s i n my  i n proportion to their the r e s e a r c h h y p o t h e s i s  f o r and a r e p r e f e r r e d  by b e a r s  marking. The e s t i m a t e d  of  (p<-05, hemlock  used  trees  hemlock, s i t k a  The f r e q u e n c y  higher  f o r a l l o f t h e marked  50 p e r c e n t o f t h e t r e e s  18 b e a r t r e e s .  western  Idid  marked by  T a b l e 4 summarizes t h e f r e q u e n c y o f s p e c i e s among  vary because  17 a m a b i l i s f i r ,  s p e c i e s r e p r e s e n t e d l e s s than  tree  by b e a r s  analysis  o f measurements  Twenty o f 21 t r e e s  (Table  trees  scarring  mean d i a m e t e r  was 23 cm dbh  the  frequency  in  the  distribution  study  o f t h e marking  (range 6.5-44.0 cm). of size  area a t the time  classes  they  t r e e s a t the time F i g u r e 15  of bear  marking  were marked.  Frequency  I observed instances  in  frequently  than  nine 1977.  o f use o f t r e e s  by b e a r s  instances of fresh  marking  I  these data  believe  that  indicate.  bears The  11  marked.  i n 1976 and rub  trees  Of t h e t r e e s  marked, 63 p e r c e n t were between 15 and 25 cm dbh when  4.3-3.  shows  trees  marks  nine more  recorded  44  Table  3:  F r e q u e n c y o f o c c u r r e n c e o f m a r k i n g by t r e e bear t r e e s i n the Ahnuhati watershed.  Amabilis other  fir  species  Z  observed  expected  17  8  1  10  (Western  hemlock, S i t k a s p r u c e , Western r e d cedar)  %  s p e c i e s among  = 18.23>%^.05  df = 1,  3.84  18  45  5-| HI UJ  oc h-  u_ O  4 3-  cc 2-  LU CO  3  1-  z 5  10  15  20  25 30  SIZE C L A S S - d b h  Figure  15:  Frequency d i s t r i b u t i o n of s i z e t r e e s at the time of marking  35  40  45  (cm)  classes of  bear  marking  46  prior  to  July  n a k e d eye.  On  bear  No.  tree  Nothing  i n the  marked  the  6 1977 June 5  were p h y s i c a l marks e a s i l y  10  and  June  leaving  sign  tree  1977,  tracks  of t h e i r in  17  in  passing  any  way.  fishing  line  July  a black  b e a r p a s s e d by  its  1977  tracks  fishing  suggested  line  I concluded their  had  that  tracks  observed  11  Between  July  fishing of t r e e s been  by  the all  of  by  1 and  the  to  of  August 5  These  distribution  the  The  km  higher  they time No.  5.  tree.  had that  5.  On  Nothing  However,  marking  in the  behaviour.  t r e e s more f r e q u e n t l y  1976  do  of  area  2  43  1977,  density  p o l y g o n i s 20  do  any  km . 2  physical  by  than  1977,  of  bear  bears.  found  the  marking  sign of  having  bears often  rub  damage.  trees  trees  16.  I  fresh  further physical  i n Figure  a r e a s but  per  of  observed  i s shown i n F i g u r e  outlined  bear  marking  suggest t h a t  bear marking  of  to J u l y  however, w h i l e e m p l o y i n g  abundance  elevation  May  seven i n s t a n c e s  not  1976  and  physical  observations  summer o f  2.2  No.  suggesting  fresh  method, I o b s e r v e d  and  this  by  substrate.  that  bear t r e e  bear tree  marking  passed  prepared  at  marked t h e  August  D i s t r i b u t i o n and  surrounding trees.  May  marking t r e e s but  spring  to  b e a r s , none o f which showed  4.3.4. The  be  was  the  would i n d i c a t e .  period  marked.  against  b e a r s may  instances  line  i t had  bears  suggested  attached  been d i s l o d g e d ,  alone  During the  that  the  It  monofilament 6  was  three  v i s i b l e to  16.  during  I  examined  I also  examined  very  marking t r e e s i n the  few area  marked enclosed  F i g u r e 16:  D i s t r i b u t i o n of bear marking t r e e s i n the watershed  Ahnuhati  4^ -4  48  Figure  16 shows t h a t t h e  confined  to  Ahnuhati  river  are trees rock  the  valley  the v a l l e y  of  bear marking  bottom  bear t r e e s a l s o  topographically of  Almost  by  of  future  similar,  km  o f major  recorded  which  the t r a p ,  t h e r e were f i v e  one  be  remove  marks and  enough  each  there  Thus, the  enough s c r a p e s and  with of  distribution that which  which  trees  a s 3.4  bear  are  in  the  marking  watercourse. observed  active  bear  behaviour  were l o c a t e d  within  trails.  had  in  association  disturbed.  no  sign  The  Ten  found t h a t  meters  vegetation the  I a l s o had  bear  nail to  a  from  in  the cover marks  imitate  several  swipes  to  produce  a  must  have  t o smooth o v e r my  a satisfactory  or  I attempted  i t required  with  n a t u r a l ground  of a track  i n the depression.  obtain  areas  bear  described  of a b e a r ' s g a i t .  surface  mark.  and  i s confined  smooth, muddy p a t c h e s a r r a n g e d  impression, suggesting that formed  While  behaviour  s c r a p e d away and discerned  of the  be  a bear e v i d e n t l y  pattern  o f moss was  The  the extreme s t e e p n e s s  density  bear t r e e s  this  trapsite  could  steep-sided.  comparison  the  Ground marking  approximate  bottom.  confined to watercourses i n areas  2 m of t h e c e n t e r of d e f i n i t e ,  I first  largely  topography.  a l l o f t h e 43  4.3.5.  valley  (Chapter 3).  F i g u r e 16 a l s o c a n  per l i n e a r  are  most o f t h e b e a r a c t i v i t y  areas  would be  purpose  polygon  the  trees  t r e e s shown i n F i g u r e 16 does n o t i n d i c a t e  restricted  For  in  i s narrow and  a r e a s mean t h a t  to  trees  water-courses  marking  on t h e s l o p e s o f t h e v a l l e y ,  cliff  are l e s s  bear  to  similar  deliberately mark t o remove  duplicate.  49  In  g e n e r a l , ground  axis  parallel  The  number  marks were e l l i p t i c a l ,  t o the d i r e c t i o n of  marks  in  a single  w h i l e as many a s 23 were n o t e d Most  marks  trapsites,  observed  fishing  of the t r a i l  sites,  marks, h o w e v e r , were n o t a s s o c i a t e d  major  from  5 t o 16,  p r o x i m i t y t o each  associated  daybeds,  their  (18-25 cm by 15 cm).  s e r i e s ranged  i n close  were  with  with  and marking  clearly  with  other.  disturbed  trees.  A few  any o t h e r  bear  bear t r a i l s  had  sign. Within  t h r e e weeks, many ground  been r e - c o l o n i z e d described  by  Martinka  were s o s p a c e d depressions two  by mosses a n d  that  and  marks a l o n g  looked  Jonkel  like  the  trail  ( p e r s . comm.).  These  I  steps  a human c o u l d c o m f o r t a b l y walk i n them.  were 2.5 t o 7.5 cm below t h e s u r r o u n d i n g  occasions  steps  found  that  ground  marks had been  soil.  The On  subsequently  r e - m a r k e d by b e a r s .  4.4.  Discussion  Marking and  common  that  both  pine  in  reflected  trees  behaviour  and ground  and,  fir.  of  the  bears  forest  t o be a  trees  available,  National composition  Park,  that  widespread  T h i s study  marked t r e e s  S c h a f f e r (1971) n o t e d  Glacier  appears  of bears i n t h e w i l d .  b l a c k and g r i z z l y  rainforest amabilis  o f both  in  the  the bears  bears  used  showed coastal  preferred lodgepole  Montana, b u t he t h o u g h t  rather than  bear p r e f e r e n c e .  this  50  The  mean h e i g h t  examined  was  communication,  139  cm.  use  o f an  would f a c i l i t a t e 631),  in their  of the  highest If  the  object  of  scent  of  marks  the  of  scent.  marking  marks  function  l o c a t e d above  dissemination  study  part  forest and  i n wolves,  I  as o l f a c t o r y  the  Peters  which  floor  Mech  (1975:  noted:  "Depositing urine w e l l above g r o u n d , on a snowbank o r t r e e , f o r example, f a c i l i t a t e s d i s p e r s a l of odor by wind, increases the e v a p o r a t i n g s u r f a c e as the u r i n e t r i c k l e s downward, and m i n i m i z e s c h a n c e s t h a t t h e mark may be c o v e r e d by snow o r washed away by r a i n . " It  i s possible that  the  t o some c h a r a c t e r i s t i c dissemination My run,  t r e e s are  trees 23,  an  marked as  more o f t e n . 73  or o n c e e v e r y density  studied. area  serves  6). as  1.4  of  high  f r e q u e n t l y as  marking t r e e . border probably  of can  d a y s on  overlap  passed through the  of  the  found  that  15  13  salmon  5 days,  and  and  marking September  area  marking t r e e s c o i n c i d e s  with  approximately  per  in their  that  km  2  use  "vocal  in  serve  to  time".  since another Mech  20  of  Perhaps  individual  suggested  wolves  that  when  other  in  transmit  (1975)  habitat  communication  f o r animals separated  may  territory,  area.  due  i n the  noted  of c o n t a c t  and  season of the  once i n e v e r y  among b e a r s  separated  Peters  be  average.  (1973)  time e l a p s e d  tell  facilitates  the  (1971)  concentration  means  their  which  b e a r m a r k i n g t r e e s was  individual  determine the  b e a r s may  b e a r s between June  o l f a c t o r y communication  between  f i r by  scent.  that, during  t i m e s by  Eudnai a  species  Schaffer  of extensive  (Chapter  whilst  The  of t h i s  indicate  were used  The  of a m a b i l i s  and/or r e t e n t i o n of  results  possibly  use  space,  messages bears  has  that,  passed at  examine s c e n t wolf  can  packs  a  the marks have  51  During  the  ground  marking  run.  During  marking  1976  field  behaviour  i n 90 f i e l d  Stonorov  and  fishing  intraspecific  at McNeil  sites.  Grizzly  olfactory  senses  fishing  bears  rely  to  unpredictable approached fishing  near  communication possibility other The  the  few  river.  intra-  high i n c i d e n c e o f g r o u n d  probably  i s related  on  same  resource.  functions  in  olfactory  functions  as  visual  the  some s i t u a t i o n s , where  space  may  in  marking  I  suggest  and we  sleeping  or  of the usual  strife  by  warning  idividuals.  n o t e d d u r i n g t h e salmon r u n bears  feeding  tree  marking  by b e a r s  while  ground  marking  that  tactile  communication  preclude  and, i n  and a u d i t o r y  cues  the  use  of  and t h e message i t c o n v e y s  may  serve  areas  r e s o u r c e s such as food.  noise  or f i s h i n g  for olfactory  conditions  water  behaviour c o u l d reduce the  communication,  substitute  comm.).  occasions,  t h e absence  of sleeping  and p e r h a p s  T h i s marking  animals  marking  pers.  grizzlies  t o t h e h i g h c o n c e n t r a t i o n of  environmental  signals.  in  fishing  with s w i f t  several  and i n t e r s p e c i f i c  of t h e presence  grizzlies  a u d i t o r y and  o f background  of  Thus,  on t h e i r (Jonkel  On  meters  s i g n a l s , ground  of both  bears  a  that  situations  ideal  associated  level  currents.  to within  ground  noted  the  apparently  cues  were f r e q u e n t l y  wind  (1972)  among  extensively  a r e a s which g e n e r a t e d b o t h a h i g h  salmon  D u r i n g t h e salmon r u n , many  f o r environmental  sites  the  was one o f f o u r  aggression  next  to  20 i n s t a n c e s o f  Stokes  River i n Alaska.  o f t h e d a y b e d s were s i t u a t e d  four instances of  days p r i o r  between s t r a n g e i n d i v i d u a l s  which r e l e a s e d  Good  I found  30 d a y s o f t h e r u n , I f o u n d  behaviour.  meetings  season,  these to  with a high r e s o u r c e c o n c e n t r a t i o n o f I t may  also  serve  to  advertise  the  52  presence marking  of  p o t e n t i a l mates.  by b e a r s  experimental  must a w a i t  manipulation.  Understanding  rigorous  field  of the f u n c t i o n of work  as  well  as  53  CHAPTER  5 - THE USE OF TRACKS IN THE IDENTIFICATION  AND  CENSUS  OF BEARS  5.1.  Introduction  The  limited  visibility  of  e c o s y s t e m s and t h e e x p e n s e o f place  a  premium  techniques of  which w i l l  systems.  One a r e a  tracks.  If  be  various areas with  individual  could  could  be o b t a i n e d .  which  leaving  similar  criteria  Further,  yield  in  both  cases  s i g n , such as b l a c k  Klein was  an  near  the  of  of  reliable  two s p e c i e s  River,  for  T h e s e w o r k e r s measured  They measured  (1959) grizzly  of the B e l l a  the r i v e r banks as w e l l as i n prepared trails.  grizzly  Edwards and Green  a tributary  would  activity.  and  on f o r e s t  species  bears,  o f t r a c k s as a census technique Atnarko  these  systems  sympatric  Columbia.  situated  of the  minimum numbers and  and g r i z z l y  technique.  their tracks  river  River, B r i t i s h mud a l o n g  by  t h a t t h e use o f t r a c k s t o census  unreliable  the value  mammal  register  i n the i n t e r p r e t a t i o n of sign of t h e i r  (1959) s t a t e d  of  o f t h e use o f  which d i s t i n g u i s h t h e t r a c k s o f these  valuable  assessed  could  i n such  t o the occurrence  P e r i o d i c monitoring  substrates  trend.  of s p e c i e s use  field  something  be i d e n t i f i e d  substrates proportion  systems  indirect,  the  h a b i t a t s and a q u a n t i t a t i v e e s t i m a t e  population  bears  in  vegetated  t o determine  potential i s  animals  then  established  appropriate  bears  of simple,  researchers  of obvious  densely telemetry  a b u n d a n c e , and a c t i v i t y  characteristics,  could  be  enable  in  sophisticated  on t h e d e v e l o p m e n t  the d i s t r i b u t i o n ,  track  mammals  tracks i n  Coola sand  substrates  14 p a r a m e t e r s  of  the  54  forepaw  impression  successive within track  prints  the  tracks-  o f t h e same b e a r  track  could  be  w h i c h d i d not  of a s i n g l e  to  show any  where b e a r s a r e s c a r c e . composite  foot  was  about  commented  that  sum  10  less  track  variation  smooth  of the t r a c k  Burton  parameters, attributable  than  measurements might  have m e r i t  (1975) f o u n d  o b t a i n e d from the  a track true  be used  that  frequency  t h e t e c h n i q u e might  measurement percent  the  irregularities  P i e k e l e k and  measured  They c o n c l u d e d  that a  f o r any  that  also  determine  and  significant  They s t a t e d  They  individual.  constructed,  individuals.  dust  124  measurements were n o t u s e f u l  curve  to  from  that  in  a  light  sum.  They  to d e r i v e  rough  e s t i m a t e s of p o p u l a t i o n composition. The bears  use in  o f t r a c k s as an i n d e x the f i e l d  (1976) f o u n d weight  because  judged  o f t h e wide r a n g e  confidence. positive  weight  and  body  predicting  weight  a  bears  weighing  a t t h e 95 Burton  the estimate  o f 48  kg.  and  kg  their  from  Best foot  of of  Pelton  and  body  in  body  data to  be  than  kg  70  percent l e v e l  (1975) f o u n d  o v e r a range  67-397 kg  50  of bears l a r g e r  California.  polar  than  composite  body w e i g h t  individual  Cherry  d e r i v e d from  of weights  between  bears i n northern  equation  smaller  However, P i e k e l e k and  correlation  measurements  the  of  between f o r e p a d w i d t h  the equations  for predicting  weight  been a t t e m p t e d .  a strong correlation  They  inadequate  caught  also  i n Tennessee b l a c k bears  weight.  black  has  to the  a strong  individual 15 t o  (1977)  180  paw  kg f o r  derived  m e a s u r e s o f 16  with a standard  of  an  wild-  error  of  55  While v a r i o u s the  tracks  1954,  of  black  Schaffer  used  of t h e  size with  identification  of  The  conducted application occurred data  for  Specific 1.  to  the were  i f  two  classes the  Green  (1959)  study  bears  the  bears  supportive  telemetry. of  bear t r a c k s  were:  characteristics from g r i z z l y  track  was  where as  of  between  adequately evaluate  used t r a c k s  by  track  potential  to a l i m i t e d area I also  of  distinction  and  d i d not  track  of b l a c k  as  of  permit  bears  characteristics  of i n d i v i d u a l bears  within  a  permit  species.  Methods  Twenty-two trapped  i f  species.  Edwards  Murie  comparison  two  and  separate  f o r example  rigorous  bears  and  of the  (see  re-evaluated  densities.  determine  separation  5.2.  I  information  determine  to  of the  l o c a t i o n s determined  separation 2.  been no  work o f  objectives  have been used t o  bears  morphology  a wide a r e a  high  bear  has  individual  of t r a c k  at  grizzly  which  previous  over  criteria  tracks  and  characteristics  species.  and  1971), t h e r e  characteristics I  subjective  track  characteristics  a n i m a l s were measured and  p a r a m e t e r s which  of  19 2  recorded.  I measured and  the  tracks  Figure  manner  in  17  and  10  displays  which  they  i n dry,  loose  recorded. Early  s a n d and  in  the  field  work, I f o u n d  i n snow were h i g h l y  tracks observed to  in these  wet  sand  be  Like  Edwards and  Green  v a r i a b l e and  substrates.  consistent  that tracks  in their  (1959), I  I subsequently  I considered  wet  ignored mud  "track-recording"  established  substrates  and  ability. which  ID Reason  Foot TL T L PW PL WA N-S T-S TP  Toej Below  Date  Jpaco  56  th  2  3  L Location  Comment  Substrate W N  Figure  17:  Parameters o f bear data sheet  t r a c k s w h i c h were measured and a s  4  5  57  would  record tracks, i n addition  areas  of s u i t a b l e s u b s t r a t e s .  5.3.  I  expected  young i n d i v i d u a l s , would be  individual of  11  a sufficient  adults.  i t  overlap  I operated  known g r i z z l y  that  was  bears  from  tracks alone.  their  are a d u l t females  and  widths on  without  therefore, bears  in  comparison very  and  = .38  be  five to  The  identical Table  separated  4  of  are  of the data  the  i n the  premise  t r a c k s to paw  known b l a c k  distinguish  first  two  broad  o f which  I  These  earlier free  of  trapped,  bias.  cm,  The  subadult  w i t h i n f o r e p a w s and  hindpaws o f t h e t h r e e f e m a l e  4).  The  had  of  and first  on t h e b a s i s o f t h e i r  t r a c k s i n the  field.  were  they  can,  two  black  deviation  the  of  researchers,  Again,  two  shows  (s)  of the  b l a c k . b e a r s o b v i o u s l y p r e c l u d e s s e p a r a t i o n of i n d i v i d u a l s species  these  w i t h i n hindpaws  and  4 at  d e v i a t i o n (s)  similarity  grizzlies  cubs  females.  (e.g. standard  Table  i n Table  measurements  data  of  individuals  different  track  found  classes  tracks  (e.g. standard 4).  separate  bears.  The  that  characteristics  three g r i z z l i e s  o c c a s i o n s , by  measurements = 0.14  on  maturity.  considered  Table  initially  not p o s s i b l e t o d i s t i n g u i s h  cm.  knowledge  widths  and  their  almost  widely  similar  forepad  of  t h a t were t r a p p e d ,  confirming  i n d i v i d u a l s are  taken  i n the t r a c k c h a r a c t e r i s t i c s  difference  possible  but i t was  forepad  in  T a b l e 4 summarizes the  individuals,  side,  which I e n c o u n t e r e d  Results  While  there  to those  of  forefemale within  58  Table 4.  C h a r a c t e r i s t i c s o f g r i z z l y and black bear paws  ( A l l parameters  measured 1n cm and d e s c r i b e d i n Figure 17).  ID  Foot  G 7P  G 21WC  G 7WC  TL  TL  N-S  N-S  16  12  13  22.-5  20.5  11  17.0  12.1  24.2  PW  PL  WATP  Space t b  th  12.7  L W N  4.2 4.0 4.2 4.2 3.7 2.8 2.9 2.6 2.8 2.5 3.7 4.5 4.9 4.8 9.0.  15.8  12  L W N  4.0 4.2 3.7 3.6 3.02.5 2.4 2.6 2.5 2.5 2.2 2.4 2.2 2/0 2.4  13.2  7.2  12.3  L W N  4.2 4.0 4.3 4.0 3.6 3.0 2.7 2.8 2.6 2.5 4.0 4.4 4.5 4.5 4.0  22.2  12.5  17.0  12.0  L W N  4.0 4.2 4.1 3.9 3.0 3.0 2.8 2.5 2.5 2.2 2.0 2.2 2.0 2.0 2.0  17.0  12.0  13.0  7.4  12.4  L W N  4.0 4.5 4.1 4.3 3.5' 3.0 3.0 2.9 3.0 2.4 4.4 5.0 5.0 5.0 4.0  23.3  21.0  12.4  16.0  11.9  L W N  3.5'4.0 4.0 4.0 3.5 3.0 2.6 2.5 2.5 2.0 2.5 3.0 2.5 2.3 2.2  L W N  6.4 6.4 5.72 5.72 5.1 3.8 3.5 3.2 3.8 3.5 5.1 6.4 6.1 5.7 5.1  L W N  5.72 5.72 5.1 4.78 3.8. 3.7 3.8 3.5 3.7 3.5 2.24 2 2 4 224 2.54 2.9 '•  Standard deviation(s)  F  0.5  0.26  0.38  0.20  0.94  H•  1.37  1.0  0.80  0.57  0.20  G 12/22  LF  19.1  13.7  18.1  7.6  17.  LH  25.1  22.6  15.9  17.5  18.4  LF  9.0  6.5  7.0  3.4  7.4  X  X  L W N  2.7 2.7 2.5 2.3 1.7 1.4 1.5 1.5 1.5 1.3 1.7 2.0 2.1 2.0 1.6  G cub o f G 21WC  "•  X  X  LH  13.0  12.0  6.5  9.3  7.0  X  X  L W N  2.2 2.3 2.3 2.0 2.0 1.4 1.5 1.4 1.3 1.2 0.9 1.0 1.0 1.0 1.0.  LF  14.5  10.5  11.0  5.2  10.2  X  X  L W N  3.1 3.1 3.0 3.0 2.8 2.0 2.2 2.1 2.2 1.9 1.5 1.5 1.5 1.8 2.0  RH  21.0  20.0  10.0  14.5  10.3  X  0  L W N  3.0 2.6 2.7 2.1 2.3 2.0 2.0 2.0 1.8 1.5 1.5 1.5 1.5 1.8 2.0  G S u n k i s t RF  19.2  14.0  12.2  7.3  12.5  L W N  4.2 3.8 3.1 2.5 3.1 5.7 4.6 4.5  L W N  4.0 4.0 3.6 3.9 2.9 -1.9 2.2 2.3 2.3 2.2 2.2 2.6 2.5 2.3 —  G 20  RH  23.5  20.2  12.7  G Sighting LF 1  16.5  11.4  13.34  G S i g h t i n g RH 2  26.7  25.4  15.9  X •> no  18.5  5.72  20.3  G =  12.7  — —  13.34  X  X  L W N  3.8 3.5 2.7 4.5 5.1 5.7 5.1 4:4  15.9  X  X  L W N  -  grizzly  yes  —  2.5 2.5 2.5 2.5 1.9  59  Table 4. (continued)  ID G Shardik  G" Sheba  B  18  B 20  •  TL  TL  Foot  N-S  N-S  PW  •PL  RF  19.0  14.1  14.0  7.9  12.7  X  X  RH  26.3  22.9 .  13.0  18.1  13.3  x  x  WATP  Space  tb  th  2  3  4  5  L u N L W N  4,.0 4 .4 4..1 4.1 3.8 2,.9 2 .5 2..7 2.5 2.5 5,.1  3.8 3.8 3.8 3.8 3.7 2.9 2.9 2.9 3.0 2.5 4.1  1,.7  2 .9 4..9 2.9 2.5 3.A  LF  17.8  13.7  12.8  6.4  12.9  X  X  L W N  LH  25.4  22.9  13.3  17.0  13.2  X  X  L W N  3.8 3.8 4. 0 3.8 3.8 2.7 3.2 2. 9 2.9 2.5 2.5  LF  11.0  8.2  9.0  4.5  10.1  0  0  L W N  2.,8 2..9 3..0 2.8 2.2 2..0 2..0 2..0 2.1 1.7 2..2 3,.0 2..8 2.7 2.0  12.2  9.3  0  0  L W N  2..7 2..5 2.,6 2.5 2.0 2.,0 1..8 1..6 1.7 1.6 2..0 1..9 2.1 1.3  LH  ....  16.7  RF  ....  9.6  9.2  4.9  10.1  0  0  L W N  3.,0 3,.0 3..2 3.0 2.0 1.,9 1..8 1..8 2.0 1.5 2..9 3..1 3..2 3.2 2.5  LH  ....  16.0  9.3 5.5  12.2  9.1  0  0  L W N  2.,7 2.1 2.9 2.5 2.0 1. 6 155 1.551.7 1.6 . 1..9 2.0 1.9 1.8 1.5  8.8'  B  19/23  LF  14.6  11.3  12.7  6.35  16.0  0  0  L W N  3..8 4.14 4.45 4.14 2.9 2..7 254 254 3.3 2 5 4 2..9 3.3 3.3 3.2 2 5 4  B  4  RF  n.o  9.1  8.2  4.6  8.7  0  X  L W N  2..5 2,.9 2..8 2.8 2.3 1..6 1..8 1..7 1.8 1.5 2..7 2,.6 2..6 2.8 2.5  RH  16.8'  15.5  8.2 4.5  11.5  8.3  0  0  L W . N  2,.2 2,.5 2.3 2.1 1.9 1..2 1..3 1.5 1.5 1.4 1..8 1,.9 1.8 1.7 1.4  B Terrace  LF  16.9  16.0  8.7  5.0  8.8  0  0  Not recorded  B Sighting  LF  10.5  7.9  9.0  4.2  9.6  0  0  Not recorded  X • no  B = black  0 = yes  60  Large from  adult  grizzly  a l l other bears  debates  bear  by  t r a c k s were r e a d i l y d i s t i n g u i s h e d  their  as t o the s p e c i f i c  large  of  characteristics known  black  the  different  separate  of both  found  black  from  had  toes while  significant  which five  the  in this  characteristics  bear  bears.  (p  =  .001)  which  were  t o e being  below  hindpaws  had f i f t h  toes  which  which had f i f t h  differed  characteristic i s t h e presence  A l l o f t h e 11 g r i z z l y  joined,  while  separate  from  a l l of each  grizzly  other bears  toes  Ten  of  (p  t o e s which  black  and  is  bears The  the  =  a  11  .004)  were below  grizzly  f o r e p a w s had t o e s  the s i x b l a c k  which  were even w i t h t h e  or absence of a space  bear  the  forepaws  There  different  i n which  second  bear  toes.  (Table 5 ) .  This i s significantly bears  of  between b l a c k and g r i z z l y  The  and  five  These c h a r a c t e r s a r e  four  (Table 5 ) .  black  and  with the m i d l i n e o f t h e other  midline o f the other toes  toes.  were  groups.  of the f i f t h  other  track characteristic  the f o u r black  paws  bears  A l l o f t h e 11 g r i z z l y were even  I  t h e 22 t r a c k  o f t h e s i x b l a c k b e a r s had f i f t h  difference  other four toes. from  compared  which  e a c h o t h e r i n t h e f o r e p a w s and  were below t h e m i d l i n e o f t h e  grizzly  two  and g r i z z l y  i n F i g u r e 18.  four toes  I  between t h e s e  illustrated fifth  frequent  o f t h e other f o u r toes as w e l l as t h e frequency  being  hindpaws  bears  size.  5 compares t h e f r e g u e n c y  midline  the toes  animal's  and  had  characteristics  o f a g r o u p o f 11 known g r i z z l y bears  significantly Table  the  We  s t a t u s o f s m a l l e r t r a c k s , however.  looked, therefore, f o r distinquishing independent  size.  between t h e which  had t o e s which  (Table  5).  in this  track characteristic  bear  difference  were were  between i s hiqhly  61  Table  5.  F r e q u e n c y o f o c c u r r e n c e o f t w o paw c h a r a c t e r i s t i c s which d i s t i n g u i s h black bears from g r i z z l y bears.  Fishers  exact  Probability Character  Grizzly  Bear Black  11  1  bear  Siegel  (1956)  F i f t h toe r e l a t i v e to middle of other four toes  fore  paws:  even below  hind  paws:  even below  Separation  fore  paws:  of  5  10  0  1  4  11  0  .001  p =  .004-  p =  .001  p =  .007  toes  joined separate  h i n d paws:  0  p =  joined separate  0 11  0  6 0 4  ure  18:  Two f e a t u r e s w h i c h d i s t i n g u i s h b l a c k f r o m g r i z z l y b e a r forepaws 1) s p a c e b e t w e e n t o e s a n d 2) t o e b e l o w m i d l i n e of o t h e r f o u r toes  63  significant  (p = . 0 0 1 ) .  A l l of the  bears  toes  were  had  different toes  which  (p = .007)  from  w h i c h were s e p a r a t e Of t h e  11  grizzly  seven  were  adults,  While  limited,  characteristic The  bears  can  be  characteristics. the  and  hindpaws o f a d u l t s and  hand, four  ( F i g . 18).  have  fifth  t o e s , and  While  characteristic,  f o r f u s i o n among t h e  s u b a d u l t s , and indicate size  toes  to  one  that of t h e  separated  bears  on  had  toes,  was  a  this  cub. track  animal.  have f i f t h  f o u r t o e s , and  juveniles.  which  are  basis  of  two  toes  which a r e  even  which a r e j o i n e d  separate  this  t h e r e a r e no  both  Black b e a r s ,  below t h e from  and  the  T h i s f e a t u r e o c c u r s on  which are  exceptions  (Table 5).  other  of the  the other  another  each  do  reliably  one  significantly  i n d i c a t e t h a t t h e t r a c k s of b l a c k  Grizzly  m i d l i n e of  is  grizzly  b e a r s , a l l of which  were  i s independent  This  the  four black  examined  data  foregoing results  grizzly  with  from  three  these  joined.  the  bears  11 h i n d p a w s o f  the  fore-  the  other  m i d l i n e of t h e  other  one  difference  occur  exceptions  in  on  to  another  ( F i g . 18).  i n the  "toe-below"  the  "toe  separate"  characteristic. It  has  also  size,  a grizzly  than  a black  the  third  forepaws data  bear  bear.  t o e , and of  been  no  overlap  cm)  and  those  is  not  independent  f o r two  would have l o n g e r T a b l e 6 shows t h e  the r a t i o  eight  p o i n t f o r the  claimed  grizzly grizzly  between of t h e  the  between bears  bear claw  cub  claws pad  on  the  width,  claw  these  and  f i v e black  i s excluded,  width,  length  which c a n  on  bears. then  This be  on the  I f the  there  grizzlies  (2.6-3.3 cm).  paw  forefeet  characters  l e n g t h s of the  black bears o f t h e pad  bears of i d e n t i c a l  is  (4.2-6.1  difference  considered  an  64  Table 6.  The r a t i o o f n a i l l e n g t h t o f o r e p a d w i d t h as a d i s t i n g u i s h i n g c h a r a c t e r i s t i c of bear t r a c k s .  The Bear  Pad  width  Claw  length  ratio  riail  length  ID  Foot  G 7P  RF  13  4.9  0.377  G 21WC  LF  13.2  4.5  0.341  LF  7.0  2.1  0.300  G 20  LF  11.0  4.2  0.382  G 7WC  LF  13.0  5.0  0.385  G 12/22  LF  18.1  6.1  0.338  Sunkist sighting  RF  12.2  4.6  Sighting  LF  13.3  5.7  G cub o f  21  (cm)  (cm)  forepad  .  to  width  0.377 0.429 X =  0.366  range 0.300  -  0.429  Bl  4  RF  8.2  2.6  0.317  Bl  19/23  LF  12.7  3.3  0.260  Bl  18  LF  9.0  2.8  0.311  Bl  20  RF  9.2  3.2  0.348  RF  8.7  3.0  0.345  Terrace  of  X  0.316  range 0.260  -  0.348  65  index to the overlap  between  grizzly black  animal's  bears  however,  that  bias t h i s among  ratio  whch  is  of claw  there  t h e low  frequency of a d u l t s  full  with comparable  because  of  12.7  widths  has  of  a claw  the  of  the  It i s possible,  growth  paws.  (25 p e r c e n t )  width  i n the b l a c k bear  differential  cm)  considerable  those  x = 0.316).  components o f b e a r  centimeter  pad  is  l e n g t h t o pad  0. 260-0.348,  (pad w i d t h = a  as  (range  the s t r u c t u r a l bear  size,  0.300-0-429, x = 0.366) and  conclusion  black  4.6  the  (range  bears  body  The  shorter  ( c l a w l e n g t h s = 4.9,  patterns  single  length  of  than 4.5,  data  adult  3.3  cm  grizzlies 5.0,  and  bears that  were  cm).  5.4.  Discussion  The  fact  examined  had  that  virtually  that  individual  basis  of t h i s  which  rely  nothinq  the  identical  bears  cannot  characteristic exclusively  11  paw be  the  grizzly  characteristics  distinguished  alone.  on  of Edwards and  grizzly  scarce.  The  bears  Green  is  Thus,  of  dry,  similarity  we  found  indicates  that  loose in  census  fraught with  the  census  of t r a c k  I concur  of  with  the  techniques produce  with  the  of t r a c k s  i m p r e s s i o n s on  combined  tracks  on  t o a r e a s where b e a r s sand  the  individual  s u r v e y s of t h e s i m p l e  error.  reliably  (1959) t h a t t h e use  sand,  indicates  measurement o f t r a c k s  restricted  high v a r i a b i l i t y  composed  would be  of  more t h a n a b s o l u t e minimum numbers.  conclusion census  three  "river  to are  bars  striking grizzlies,  drift"  type  66  It  i s possible  distinguishing impression.  that  characteristic All  of  drawbacks,  however.  absent  their  from  reliable  the  While  individuals,  for infection  unlikely  that  i t caused  removal.  Again,  fill  the  w i t h mud  that  the  use  technique  with  Track  and  would  of  are  combined  the have  be  a  would s e r v e  to  in  death.  would  incurred to  as  a  s i n c e such I concur  acting It  is  s i n c e the  result  infection.  useful  of  would p e r s i s t  to distinguish  census  in  considered  probably  heal eventually .  tracks  limited  paw  damage  a  impressions are f r e q u e n t l y  probably  be  I  result  to the  would n o t  register  serious limitation  which c o u l d  t h e bear  marked w i t h  o f t h e n a i l s would n o t  the  in  its  result  of  Branding  or  a mark c o u l d with  individual  others  bears  is a  application.  measurements a r e u s e f u l ,  impressions telemetry  i t has  tissue  pad  which  claw  would e x p o s e t h e a n i m a l  of  would  toe c l i p p i n g  m a t e r i a l added  aggravation  cutting  methods  tracks, clipping  as a s i t e  attachment  b e a r s c o u l d be  which  Since bear  technique.  distinguish  individual  with  l o c a t i o n s or automatic  however, when other  perfect  information  track  such  camera r e c o r d i n g s y s t e m s  as  (Smith  1978). Tracks  were f o u n d  t o be  and  grizzly  bears  Of  t h e two  distinguishing  "space"  latter  when p e r f e c t  c h a r a c t e r t o be  characteristic,  useful  as  characteristics  from  reliable  are  character objectively.  length-of-claws  track impressions  more  there  f o r the s e p a r a t i o n  the t o e s  found,  than  difficulties On  were  larger  were  the in  bears,  reliable.  of  black  available.  I consider "toe  the  below"  a s s e s s i n g the size  and  the  67  CHAPTER 6 - FOOD HABITS  6.1.  Introduction  One of  of the b a s i c p r e r e q u i s i t e s to understanding  a mammal s p e c i e s i s  habits.  Many  aspects  as i t s d i s t r i b u t i o n understood  an  fully  accurate  of a species l i f e  when t h e y  used by t h e a n i m a l .  food  species  consumed  than  a  their  omnivorous nature  for  century.  i s needed n o t o n l y  a l s o on t h e i r  (Mundy  1963, Hamer  a l . 1977,  Food  1972), t h e i n t e r i o r  (Murie  Island  ( C l a r k 1957), and t h e A l a s k a n  well  of  the  1972, H a g l u n d  black  very s i m i l a r t o those food  Glacier  1977),  Montana  peninsula  habits  of  bear  1968,  and  of the g r i z z l y  the b l a c k bear  bear.  (Hamer  mountains o f  1978),  (Chatelain  Kodiak 1950),  ( s e e f o r example  Kistchinski  h a s documented f o o d  1975),  Columbia  Alberta  (Jonkel  been  National  of British  a s s e v e r a l a r e a s i n E u r o p e and R u s s i a  Zunino and Herrero Study  northern  have  (Pearson  Nagy and R u s s e l l 1978), t h e i n t e r i o r  Alaska  as  1975),  associated  bears  Yukon T e r r i t o r y  (Mealey  f o r more  our concept o f  variations  habits of grizzly  1974, L l o y d and F l e c k  1944),  importance,  known  have r e f i n e d  documented  southwestern  on t h e  and a b u n d a n c e .  studies  and  (Martinka  relative  have been b r o a d l y  N a t i o n a l Park  Montana  et  Recent  f o r the  Park,  (such  Information but  geographical areas.  Yellowstone  strategy  to the p r o p e r t i e s of the  food h a b i t s o f bears  assessed  i t s food  are related  n u t r i t i o n a l value, d i s t r i b u t i o n ,  with  history  of  and abundance i n s p a c e and t i m e ) c a n o n l y be  foods  The  description  the ecology  1972).  h a b i t s which a r e  Among  other  have been d e t e r m i n e d  areas,  i n Montana  68  (Tisch  1961),  Washington Pelton  interior  (Poelker  (1977)  in  the  black  home  r a n g e , and  grizzly  have n o t  Poelker  and  Atwell et  of the  Hartwell  study  fall,  to the study  of  1.  to  food  bears  identify  to r e l a t e  the  understanding described There the  are  seasonal t o be  grizzly  bears,  The  for  coastal  and  bear-human  and  Chatelain 1974,  North large  1950,  bears  American geographic  Clark  Berns e t a l .  c o n s t r a i n e d by  sample e a r l y i n t h e  1957,  1977,  and  o f Smith  required to  food  h a b i t ' s study  late travel  use  of food  by  were: black  relationships  food  to i t s use two  aspects  times.  seasonal  by  of bear  of  bears,  preceding  quality  and  and  and  objectives  feeding ecology  to  current  as i t i s  literature.  potential  limitations  presented  reliable  and  effort  other  s p r i n g , nor  Columbia,  of food  any  I have had  data  Beeraan  habits.  incomplete  (see  coastal  of b l a c k  in coastal British  i n the  two  use  appear  area,  are  o b j e c t i v e s of the  availability 3.  food  h a b i t s was  at those  g u a n t i f y the  to  productivity,  p r i m a r i l y because of the  grizzly 2.  Recently,  movement p a t t e r n s  Russell  I d i d not  area  Specific  1973,  and  1977).  project.  the  bears  1972),  1973).  the  h a b i t s data  been s t u d i e d  al.  This  in  attributed  a v a i l a b l e food and  (Hatler  Hartwell  Smoky M o u n t a i n s t o t h e i r  The  areas  and  have  interactions,  Alaska  here.  to the First,  method t o s e p a r a t e  s i n c e the  species  t o assume t h a t t h e i r  d e s c r i p t i o n of t h e r e does  s c a t s of  were s y m p a t r i c food  habits  black  i n the  were  (1978) i n d i c a t e t h a t t h i s a s s u m p t i o n  not and  study  similar. is  valid.  69  The  data  foods not  presented  used  by b o t h  possible  used.  the  o f a combined  sufficient  observed  shifts  using the  of  the observed  10 p e r c e n t  from  1977.  collected  I  scat  been  possible  information  to  presented  o f the  Second, i t i s  samples  to  treat  i n the p r o p o r t i o n s of  method  of  Hanson  volumes a c t u a l l y  the t r u e v a l u e s f o r the nine  sample  and b l a c k b e a r s .  72 s c a t s would h a v e been n e c e s s a r y  percent  have  bears  collect  F o r example,  (1956),  consist  grizzly  to  statistically  thus  and  Graybill  t o be c e r t a i n differed  sample  foods  that  by l e s s  period  of  95  than May  scats f o r this  p e r i o d and i t would n o t  collect  than  is  more  limited  to  a  30.  Thus,  sketch of the  the actual  situation.  6.2.  Methods  6.2.1.  Scat  collection  The c o l l e c t i o n ,  analysis,  d a t a f o l l o w s t h e methods collected within  fresh  bear  the study  within  a week o f t h e i r  of  of Tisch droppings  a month o f c o l l e c t i o n .  through  and p r e s e n t a t i o n  t h e s c a t were r e c o r d e d  with a f e l t  bags.  habits  their  We  contents  we s y s t e m a t i c a l l y t r a v e l l e d  c o u l d be  deposition.  food  and H u s s e l l ( 1 9 7 1 ) .  and d e t e r m i n e d  Since  area, droppings  (Trademark) c o l l e c t i o n  (1961)  of  aged  and  collected  The d a t e , a g e , and pen  on  plastic,  location Ziploc  70  6.2.2. The  Scat  food  (1972) f o u n d relative  analysis  habits that  material.  the To  enamel  dissection  fishing  line.  components  scats.  a  index  reliable  were  were  evenly,  excess  sieve. tray  water,  The s t r a i n e d gridded  Ten s u b s a m p l e s  predetermined :  is  by e x a m i n i n g  Hatler to  the  of most f o o d s by b e a r s .  locations  Subsamples  points  analysis  remove  through a k i t c h e n  grid  studied  t o a n a l y s i s , d r o p p i n g s were s o a k e d  them, t o mix  to  scat  consumption  Prior  were  placed  points  in  on  identified  and  to  the  scats  s c a t was  with  from  from  a  a  Petri  i t .  Plant  remove  monofilament  random  according  number which  an  eight  the  study  area  power  table. had  fragments l y i n g  with  strained  s p r e a d o u t i n an  the t r a y  dish  soluble  were  moveable  were t a k e n  generated  i n water t o s o f t e n  on  10 these  dissecting  microscope. Plant comparison  samples with  Identifying  from  the plant  fragments observed  characteristics  photographed  were  and a r e r e c o r d e d  of  the  in  plant  collected the  droppings.  fragments  i n a separate report  for  (Fleck  were eta l .  1977). The  f o r m u l a e u s e d t o c a l c u l a t e volume  importance  value  ( 3 ) , and r e l a t i v e  (1),  importance  frequency value  (2),  (4) a r e a s  follows: Volume  (%) =  Frequency  £ volumes of food  (%) =  Importance Value  item/# o f s c a t s  x 100  E # scats  with food  (IV) =  % volume x % f r e q u e n c y / 1 0 0  I  item/* of s c a t s  (1) x 100  (2) (3)  71  Percent  Importance  Value =  £  IV o f one f o o d  item/  IV (4)  of a l l food  6.2.3.  Bear f o o d  Representative were  collection  and s t o r e d  The  regimes: and  and a s p e c t .  At  content each  bear f o o d s .  samples  from  a t 50°C w i t h i n  sites four  A l l sites  were  A line  was  samples  were  light  by  moisture canopy, i n their  were c o l l e c t e d variation  Nutrient  on in  b a g s and  collected:  moisture  represented  this  transferred  a  line. to  the site The  duffel  t o U.B.C.  were e s t i m a t e d f o r e a c h o f t h e f o o d content,  (ADF) , c r u d e p r o t e i n ,  50°C.  along  of the  analyses  samples  and g r o s s e n e r g y .  (wet) samples  and m o i s t u r e w i t h i n  collected  in plastic  nutrients  at  and  to reduce p o s s i b l e  180 t o 250 g  i n available  following  weight  explored  identical  c h o s e n whose e n d p o i n t s  The  fibre  the  4 days o f  wet-open  samples  bags w h i c h were k e p t c o o l d u r i n g t r a n s i t  crude  was  light  canopy,  shower  in  analysis.  dry-closed  I collected  were p l a c e d  6.2-4.  identified  due t o l e a c h i n g .  site  composite  dried  Where p o s s i b l e ,  a rain  maximum d i f f e r e n c e and  foods  between  samples  canopy.  t h e day f o l l o w i n g nutrient  and  of variation  plant  wet-closed  bear  f o r future nutrient  dry-open canopy,  elevation  of  collected  magnitude  replicating  collection  samples  droppings  i t e m s x 100  A l l samples With  the  total  ash,  crude f a t ,  s o l u b l e c a r b o h y d r a t e s (NFE) ,  were i n i t i a l l y  dried  exception  samples  of  to  constant used  for  72  determination sample  material  analysis of  of m o i s t u r e  was  was  not  any  was  (ADF)  method  Soest  (1971);  Crude  technique The and  of Van  (AOAC  furnace  The  ml  20  with  Seasonal  the  of  ladyfern,  60  (1967) a s m o d i f i e d  was  estimated  with  Mg,  t o ash  resultant  and  at  ash  This solution  water and  use  by  Waldern  the G o l d f i s c h  K i n the from  a  475° C i n a  was  was  roots one  muffle  dissolved in then  7.5  diluted  stored i n a polyethylene with  a  to  bottle Varian-  Spectrophotometer.  of  and  the  diet  s p i n y wood and  food  from  percent  May  through  September i n 1976  vegetable  matter,  25  percent  t r a c e q u a n t i t i e s of i n s e c t s .  a r e a consumed  insects,  consisted  techniques  acid-detergent-fibre  Wine  burned  The  collected  salmon,  study  club,  by  e l e m e n t s Ca,  was  HCL.  distilled  averaged  bulk  sample  percent  Droppings  the  nutrient  Results  6.3.1.  percent  energy;  the f o l l o w i n q e x c e p t i o n s :  c a b b a g e were d e t e r m i n e d  Atomic A b s o r p t i o n  6.3.  The  standard  t h e c a t i o n c o n c e n t r a t i o n s were d e t e r m i n e d  Techtron  1977  skunk  f o r f o u r hours.  of hot,  gross  1970).  l e a v e s of  gram s a m p l e .  until  content  c o n c e n t r a t i o n of the  i n the  100  fat  and  further.  with  determined and  ash,  f o l l o w e d the  p r o x i m a t e a n a l y s i s (AOAC 1970) f i b r e content  total  dried  done a t U.B.C. and  crude  ml  content,  21  came  different from  fern, salmon.  o f s e d g e , h e r b s and  eight  of  huckleberry, The  these  and  foods:  the  early  in The  sedge, devil's  summer  current  15  bears  foods.  salmonberry,  s p r i n g and  ferns,  fruit,  The  recognizable  and  diet  year's  73  growth  of  shrubs.  August and  Fruits  importance  19 and  20 i l l u s t r a t e  value  o f major f o o d s .  mean volume, i m p o r t a n c e food  grizzly  spring  items bears  The  use and  (July to  of food  early  IV),  percent  leaves  pattern  example, the  the  (May  frequency  relative  droppings  are  in of  occurrence,  importance of  relative  Ahnuhati  value  of  black  and  shown i n A p p e n d i x I I I .  to June),  early  period, (33.3  IV),  of  and  major p e r i o d s : (2)  (1)  summer and  fall  major  summer  the  percent (3)  of  spiny  foods sedge snow  diet.  bears  was  composed o f f i v e  IV),  (2)  shoots  (3.2  vegetation.  receded  fern and  The  percent  In  sedge  of  club  salmonberry (19.9  percent IV)  (App.  bears  thus  percent  IV),  and  III).  The  consisted  r e c e s s i o n o f snow c o v e r  spring dictated  were n o t  increase i n the  (15.2  grizzly  used f i r s t .  and  of d e v i l ' s  wood f e r n  huckleberry  green  diet  leaves  o f c o a s t a l b l a c k and  corresponding in  1977  of g r e e n u p i n t h e e a r l y  of those  The  changes  be d i v i d e d i n t o two  S p r i n g and  this  spring diet  As  can  summer  (1) s e d g e  entirely  i n the  and  (4) l a d y f e r n and  (5)  extensively in  September) .  During  (25.9  seasonal  v a l u e , and  identified i n 1976  6.3.1.1.  foods:  salmon were u s e d  September.  Figures  all  and  late  April  the  availability  of  1976,  available  to foraging  developed,  there  p r o p o r t i o n of v a l l e y  and  for bears.  was  bottom  a  sedge  74  F i g u r e 19:  S e a s o n a l s h i f t s i n the use o f f o o d by b l a c k and b e a r s i n t h e A h n u h a t i s t u d y a r e a , 1976  f^nj  l a d y f e r n and s p i n y wood  ^  sedge shoots of  W  fern  salmonberry  shoots of d e v i l ' s  club  l e a v e s o f t a l l b l u e h u c k l e b e r r y and alaskan huckleberry b e r r i e s o f t a l l b l u e h u c k l e b e r r y and alaskan huckleberry b e r r i e s of salmonberry berries of d e v i l ' s skunk  cabbage  salmon unidentified  club  grizzly  Percent  Importance  Value  76  F i g u r e 20:  S e a s o n a l s h i f t s i n the use o f f o o d bears i n the Ahnuhati study a r e a ,  ladyfern  by b l a c k a n d 1977  a n d s p i n y wood  fern  sedge shoots  of  salmonberry  shoots  of  devil's  leaves  of  tall  club  blue  huckleberry  alaskan huckleberry berries of t a l l blue huckleberry alaskan huckleberry berries of salmonberry berries  of  devil's  berries  of  stink  skunk  cabbage  salmon sand and  gravel  unidentified  club currant  and and  grizzly  78  When l a d y f e r n in  patches  ground  in  and s p i n y wood f e r n a  curled  form  portion o f the plant  develops  first  develop,  (fiddleheads).  Bears  i s consumed a t t h i s  time.  above  The  time  fern  the bears  20 cm o f t h e p l a n t .  consume t h e newly  during spring  occur  The e n t i r e  and u n f o l d s by t h e end o f J u n e a t w h i c h  remove o n l y t h e u p p e r  they  and e a r l y  developing  summer.  shoots  of  These occur w i t h i n  salmonberry 50 cm o f t h e  base o f t h e s t a l k . D e v i l ' s c l u b begins i t s annual of  new  leaf  on  this  plant  the and  by b e a r s i s v e r y v i s i b l e .  diet  club  of  root  The a n i m a l s  pull  stems, which may be a s h i g h a s t h r e e  skunk  bears,  of the f i r s t  Bears  Sign of f e e d i n g down  metres,  o f f the green t o p .  Although  one  material at the top of the stalk.  devil's bite  d e v e l o p m e n t w i t h an u n f o l d i n g  dig  up  c a b b a g e i s n o t a major  food  in  the  annual  s i g n o f f e e d i n g on t h i s f o o d i s common. plants i n this  plant  above t h e r h i z o m e s  the  herbaceous  cover  a n d consume t h e w h i t e  and d i s c a r d  the  green  to  It is  develop.  portion  portion  of the of  the  plant.  6.3.1.2.  During the diet,  this  and f a l l  period,  17 d i f f e r e n t  12 o f w h i c h o c c u r a t  (App. I I I ) . which  Summer  are  huckleberry (3) b e r r i e s  Of t h e s e qreater (24.4  greater  f o o d s were r e c o g n i z e d i n than  trace  12, n i n e o c c u r a t i m p o r t a n c e  than percent  of salmonberry  three IV),  percent: (2) salmon  (10.8 p e r c e n t IV) ,  (1)  quantities value  levels  berries  of  (20.1 p e r c e n t I V ) , (4) l a d y f e r n and  79  spiny  wood f e r n  (6)  berries  Thus, bears  i n the  study to  the  of c a p t u r e d  tail,  and  them.  carried  found  area  sedge  use  initial  As  (1961)  them  daybeds o f b e a r s  which  and  in late  August  two  had  many  salmon  period;  covers,  bears  became  consuming  report  thickets.  1977  gill  salmon a t t h e  adjacent  The  consumed  without  (1971)  green  bears  progressed,  Gard  of  the  ( F i g . 20).  the  salmon  IV).  river,  month  of (4.8  percent  salmon r u n ,  killed  club  salmon.  a  IV),  berries  diet  covered  e i t h e r consumed into  spring  season  and  off  (3.9  e x c e p t i o n of  the  occasionally  Bears  a  p a r t of the  sacs.  (7)  salmon e n t e r the  of b e r r i e s  ending  salmon with t h e  and  the  of b e r r i e s  percent  leaves of d e v i l ' s  from  diet  (8.9  percent IV),  (8)  and  switch  J u n e and  Meehan  observations.  (5.5  percent I V ) ,  summer  sperm  more s e l e c t i v e  or  a  in late  During  (5)  (9) l e a v e s o f s a l m o n b e r r y  show t h a t t h e  beginning  all  IV),  the b e r r i e s r i p e n  vegetation data  (5.5  I V ) , and as  percent  of d e v i l ' s club  stink currant percent  (8-9  similar  capture I  site  occasionally  carcasses  with  them. While of  the  bears,  we  salmon. bears.  dense v e g e t a t i o n g e n e r a l l y p r e v e n t e d  were a b l e t o o b s e r v e  This  provided  In t h e f i r s t  kg)  was  The  bear  observed had  dragging  On  black  which d i s a p p e a r e d  it  was  small  second  sighted next,  size  into  o f the f i r s t  black  from  feeding  bear  (8 kg)  salmon  o c c a s i o n , B.  i t was bear  the  bears  into  (approx.  on  50  sand-bar.  the  adjacent  Smith f o l l o w e d an  view f o r 20  to  a  seconds  and  h o l d i n g a salmon i n i t s j a w s . and  on  t h e s u p p l y o f salmon  w i t h a l a r g e chum s a l m o n  thicket. bear  black  instance, a small  difficulty the  insight  two  observation  t h e wide d i s p e r s i o n of  adult when The  spawning  80  habitat  in  the  Ahnuhati  available  t o any  with  the second  which  which  pursued  size  suggests  that  spawning  of bear i n the v a l l e y . bear c a p t u r e d salmon,  The  and  salmon  are  apparent  ease  the  salmon e x p e r i e n c e d i n s h a l l o w water  difficulty support  this  conclusion. Frame  (1974) commented  probably large and  decreased  the  common  in  the  efficiency led  to  water  of  greater  feeding  Periodic  flood  decreased.  The  floods  because  the water  levels  dropped,  they  streams  salmon f r e q u e n t l y  the  downstream  floods  the  efficiency  Creek  of b e a r s s i n c e  during  salmon a c c e s s t o s m a l l s i d e c h a n n e l s and As  Olsen  b o t h washed t h e s a l m o n  and  probably  floods of  efficiency  bears.  Ahnuhati,  of bears  periodic  fishing  flow of s i l t - l a d e n  h i d them f r o m v i e w  that  were the  also  fishing  ultimately  permitted the  of  the  river.  became t r a p p e d i n  these channels. During  1976,  Federal Fisheries  and  chum s a l m o n e n t e r e d t h e A h n u h a t i  The  chum s a l m o n began s p a w n i n g on  e n t e r t h e spawning h a b i t a t salmon one  was  fifth  size  record  the size  size.  of the  two  The  1976  there  salmon  records  are several  (Meehan 1961, of  bears  Russell  21  1976  feeding  on s a l m o n e g g s i n t h e g r a v e l beds o f t h e at  this  time  I collected  exposed  run  of  less  than  duration,  and  there salmon  On  levels  This  of bear p r e d a t i o n  1974),  eggs d e p o s i t e d by  22,  p e r s . comm.).  (1974).  studies  Frame  pink  the p i n k s d i d not  timing,  f e e d i n g on and  100,000  s a l m o n r u n was  The  existing 1971,  1 and  weeks l a t e r .  1977  run.  (Darke,  redds.  water  October  Gard  River  August  o f p r e v i o u s r u n s i s g i v e n by While  on  of  until  estimated that  are in  evidence of Ahnuhati.  redds formed  by  no  gravel bears Low  spawning  81  salmon i n August the bar  gravel  in  and  85  volume. bears  to the a i r - w a t e r i n t e r f a c e Although  The  gravel  large  not  able  surrounding droppings Mealey  to  of e i g h t  and  (1975) f o u n d  grizzly  droppings, the  contents  the  15  percent  the  0.11  pdm  crude f i b r e ,  70.3  percent.  food  v a l u e , but During  feeding  fact  1976  (pdm)  sign  no  12.0  pdm  observed  I d i d not  n e s t s were i n t h e ground frequently  eggs  t o o b t a i n from a  single  nests.  the bears  the  3 bee  or i n f a l l e n  wood.  associated  b e a r s on  to  pdm  with  more t h a n one  detected the n e s t s .  the  bears.  eggs  to  4.8  pdm  digestibility  gravel  bars. of  bears  I observed  nests.  A l l of  the Some  nests r o o t s of of  occasion.  of  potential  1977  The  be  crude f a t ,  instance  In  wasp and  nests contained developinq larvae.  how  from  are thus of s i g n i f i c a n t  observe  24  14.6  a minimum a p p a r e n t  Several  clear  but  effectively  protein,  were  by  beds,  e g g s were d e t e c t e d i n  ground  disturbed  river  s o l u b l e c a r b o h y d r a t e , and  be d i f f i c u l t  o f b e a r s on  were  the  c o n t e n t of t r o u t  crude  on wasps o r bees a t t h e i r  feeding  eggs  that  proximate  Salmon may  the  salmon  t h a t t h e eggs a r e v e r y d i g e s t i b l e  p e r c e n t dry matter  m i n e r a l s . . He  black  two  amount o f g r a v e l consumed s u g g e s t s t h a t  68.5  total  d i d observe  by  The  suggests  any  flesh  isolate  gravel.  gravel  and  were a b l e t o d e t e c t salmon e g g s i n  were  of the  in  redds.  examination  percent  levels  I d i d not observe  s e v e r a l s e t s of t r a c k s of both  cursory  averaged  water  a c t of f e e d i n g at these redds, I  bears at t h e d i s t u r b e d From  Interstitial  of e x a m i n a t i o n .  the  grizzlies  September.  were c l o s e  at t h e time  bears  and  the  the  these  in  the  trees. nests  I t was  not  82  6.3.2.  Nutrient  6.3-2-1. The crude  Overview  shown  i n the study in  the  sites  The  actual  there  variation  The  fibre,  and  21  gross  m o i s t u r e regimes  p r o t e i n , crude decrease  salmonberry,  i n Appendix  constant  devil's  crude (Figs.  club,  As  than  and  i n t h e p a r t which  the b e r r i e s  fibre  c a r b o h y d r a t e s , and  and  IV and,  The  food  levels taken  Methods). i n general,  number  of  sites  a l l s p e c i e s o c c u r r e d at  within  species occasionally  T y p i c a l l y , however,  the  sites.  carbohydrates, fibre 21,  the  and  increases; 23,  25,  and and  huckleberry, the of the plant  levels  of  of  moisture energy  and  26).  For  fibre:protein  which  i s n o t consumed  mature, t h e l e v e l s  decrease,  (see  and  bear f o o d s mature, the l e v e l s  f a t , soluble  while  bear  The  a r e t h e mean o f f o u r  i s much l o w e r i n t h a t p a r t  consumes  sites.  sample l o s s .  'green'  ratio,  f o r each 26).  energy,  mean v a l u e s o f t h e s a m p l e s  between  i n the f i g u r e s  ash r e m a i n  through  number o f s a m p l e s  g e n e r a l , as t h e  content  (Figs.  date  s p e c i e s s i n c e not  o f l a b e l and  plotted  In  crude  crude  collection  values are tabulated  v a r y w i t h each  vary because values  area  of v a r y i n g l i g h t  sites.  24).  protein,  f i g u r e s are the  is little  sampled  ratio  Foods  results  c o n t e n t a r e shown by  from  total  o f Bear  f a t , s o l u b l e carbohydrates, ash, f i b r e : p r o t e i n  sampled  crude  of  l e v e l s of crude  moisture  all  Characteristics  the  (Figs.  crude  animal 20,  protein  22, and  of c r u d e f a t , s o l u b l e  moisture content increase  (Figs.  21 and  25) .  83  E Z  UJ  h=20"i O 0C  UJ  40-  Q--IO-J  UJ Q  20-  UJ Q  Q- —  £ O  B-  ^60-] •  ?-30-  e  •  B  •  O  S  A  M  M  E •a 6 H a.  6H o  <  <  UJ Q  o  ^ 2H  2  D  13  e  s E  4H  J  A  -O-  G  O  60-  •I  "  s  20-  Q.40UJ u.  z 20-  10H  a- -  D-  UJ  A  M  2»  &20-I O  O S  6. 10H  H Z  80H 60-I o 40-  O  Figure  21:  Seasonal s h i f t s b e r r y (Rubus  new A  i n the n u t r i e n t speatabilis)  percent  dry  concentrations  o  berries  matter  •e  A  •  shoots  edible portion of p l a n t pdm  ©.  e  S  A  s  20-  UJ UJ  I  J  § &100-  H  •6=  —  of  salmon-  leaves stalk inedible portion of plant  84  E  xs  E  P-30-  ^60• UJ 40-  UJ  h= 20O  EQ  e a, 10UJ  o  UJ  ~T~ J  £E  A  JL  ^  JL  A  J  JL  A  J  JL  A  6H  UJ  2H  EC r JL  O  M  A  60-  J  •I 20Q.  Q.40I  "X.  UJ  < 10-  y_ _ _  20-  z  I  J  UJ  I  BE  JL  Sioo-,  g20-,  O 80 2 60-|  o  £  EC UJ CO  UJ  O  M 22:  A  o  J  Figure  JL  4H  n  E  O  E •o a •  O ^ < o  20-  Q  A  JL  EE UJ  a,  ~> 2040  M  Seasonal s h i f t s i n the n u t r i e n t c o n c e n t r a t i o n s of combined s a m p l e s o f l a d y f e r n {Athyvium filix-femina) a n d s p i n y wood f e r n (Dryopteris austriaoa) frond pdm  percent  edible portion of plant dry  matter  85 E -o  I  Q-30-  Q. •  LU  CC UJ  [Z20-  O  CQ  ee  a-10-  UJ Q  •  •  UJ Q  A  ce o  H  3  CC  O  <  •  B  40-  20 H  3  E a  6H  >  60-  J  A  Q-  Q-  •  1  1  1  J  A  6-  r-  4-  4-  2  O  UJ  ^ 2-  a CC  o  A 60H  E •o 20-  E  a. •  E.40UJ  2:  u. 20-  UJ CC  ~r  60-  S  S, 10  Z  4  0  £0  U.  fl£ UJ  A  a.  S e a s o n a l s h i f t s i n the n u t r i e n t c l u b {Oplopanax horvidum) < §|gj  new  edible portion of p l a n t pdm  peVcent  dry  matter  B- ^  •  —© ~~ •&  -  |  |  a  I A  J  fvl  concentrations  •  shoots  (St  1  "  UJ 20O  UJ  23:  100-1  O 80-  o  1  M  3 £  A  • 20-1  Figure  1<H  <  of  devil's  stalk inedible portion of p l a n t  86  E T3  «<3oH  S  40-|  O  eg u. 20-  Q-10  UJ Q  cc LU Q  T  J  cc  JL  A  S  £ O  O  E a  (3  i H <  < o  *  E  JL  A  JL  A  JL  A  JL  A  I S  6H  LU  2-4  DC  O  T" M  JL  A  60  E •o o.  Q-40H LU  20-  X  (JL  ioH  w  Z20-  ~r  J  JL  LU £E  A  LU20-|  S 6 5  o  &. ioH \  1  0  O  I  LU  E  A  JL  0  40-  LU  m  n " 600  8  t z  EC LU  24:  60H  a  z  m H-20-)  Figure  "I  LU  20-  T  a.  Seasonal s h i f t s i n the n u t r i e n t c o n c e n t r a t i o n s of c a b b a g e {Lysichitijm amevioanum) ©  L_j  lower  stalk  edible portion of plant pdm  percent  dry  matter  •  skunk  leaves inedible portion of plant  T S  87 E  T3  Q.30-  I 1=  £20O  CC  a. 1<H  ~r A  J  UJ Q  3  CC  O O  —A  J  0  4-  <  O  2-  *  A  6CH E  •§  Q.40-  1 x  UJ  T J  UJ  EC 3  A  A  iVl  H100-  • °2  H  o  O  8  °-  S 60-  ct 10-  H Z UJ  CC UJ GQ  a  40H  20  CC  A 25:  10H  CO  ^20-  Figure  20-  a  A  UJ fiL  Seasonal s h i f t s i n the n u t r i e n t c o n c e n t r a t i o n s s a m p l e s o f t a l l b l u e h u c k l e b e r r y {Vaccinivm a n d a l a s k a n h u c k l e b e r r y (v. alaskaense) §H A  new  leaves  shoots  berries edible portion of p l a n t  pdm  of combined ovalifoliwn)  percent  dry  matter  inedible portion of p l a n t  88 E  E  -a °-3<H  Q. 60  •  SS  CC UJ 4 0 H 63  £20H  O tr  Q  UJ  Q  M  3 CC  n—:—i  J  o  JL  r  A  6-  -®-  4O *  20-H  UJ  a. 1 0H  ®  3 CC  O  M  E •a a i H if  4  Q  2  3  — ©  CC  1  M  1  J  1  JL  r  o  r  J  A  60H  A  6H  UJ  2-  JL  JL  A  JL  A  JL  A  E  E  •5 20-  Q.40H  a w <  •  UJ  20-  10H -©-  T J 5  2  0  JL  A  S  ^  100  ->  O ©S 60H K- 40-  a. i o H CC UJ  0  •o-  UJ 20O  M 26:  M 8  o  Figure  UJ CC 3  Seasonal  JL shifts  (Carex s p . )  A  i n the nutrient  leaf pdm  percent  CC UJ &,  concentrations  of  sedge  edible portion of plant dry matter  89  6.3.2.2.  Nutrient content  o f bear  I sampled  three parts  salmonberry  content; and  the  during  t h e s h o o t s and berries.  Hay  probably decrease  higher  strongly  per  in  in  the  that  spring.  The  constancy a l l  and  the  develop  on t h e s a l m o n b e r r y  pdm  t h e s e a r e consumed by not  consume  These p a r t s o f t h e  plant  soluble  carbohydrates  d u r i n g J u n e and shoots  ( F i g . 21).  constituted may stalk shows (Fig.  August  require  a f a r lower a  of s a l m o n b e r r y that 21) .  the  (Figs.  20  for  as are  to  25)  sample  of  the  f l o w e r s of  protein, crude  shoots  Bears the  of  21  that  salmonberry. food  crude  fibre  values  l e a v e s and  expected, very  August  levels  on  fat,  i s lower for  the  flowers obviously  biomass than t h e s h o o t s and,  and,  stalks  energy  corresponding  higher handling time.  is  of the gross  S c a t a n a l y s e s showed  of c r u d e  the  and  levels  protein  a r e h i g h e r and  than the  However,  (pdm)  r e p r e s e n t a more n u t r i t i o u s  a p e r gram b a s i s s i n c e t h e l e v e l s and  salmonberry  f o r the August  with crude  bears.  flowers,  I n September, new  t h e l e a v e s and  p l a n t may  nutrient  protein  v a l u e o f 6.5  o f skunk c a b b a g e a r e i n e r r o r .  did  of  Crude  foods  t h e v a l u e o f 5.5  sample  bears  shoots  the  gram f o r  t h e s h o o t s of s a l m o n b e r r y  and  their  l e a v e s , o t h e r l e a v e s and  i n August.  kcals  suggest  for  t o 25 p e r c e n t o f t h e d r y m a t t e r  earlier  10 pdm  v a l u e s a t 4.5  their  Crude p r o t e i n  i s close  to  of  foods  therefore,  do n o t consume t h e nutrient  f i b r o u s and  low  analysis  i n nutrients  90  Mature s a l m o n b e r r i e s levels  (19-25  pdm) , a n d Their  pdm),  high  high  have  high  moisture  net  comparatively  low  s o l u b l e , carbohydrate content  (84-86  value  probably  nutrient  crude  fibre  levels  (46-58  percent)  ( F i g . 21.) .  o f f s e t s the c o s t s of  handling. Ladyfern in  the study  fern is  and s p i n y wood f e r n a r e t h e two f e r n s used by  are  area,  although  a l s o common t h r o u g h o u t  the highest  which I examined sedge  are  (35 pdm)  interesting  although  reflection  of  in  their  (Figs.  the f a l l their  and  and sword f e r n  and s p i n y  of  oak  the  wood f e r n  spring  foods  ( F i g . 2 2 ) . T h e s e two f e r n s a s w e l l a s low f i b r e : p r o t e i n  22 and 2 6 ) .  which appear i n t h e d i e t  sampled,  fern,  area,  Ladyfern  crude p r o t e i n l e v e l s  throughout t h e season food  bracken  the study  common on t h e most x e r i c s i t e s .  exhibit  a  deer f e r n ,  bears  of bears  levels  (< 2)  The f e r n s a r e t h e o n l y  throughout  a r e low.  relatively  ratio  the  period  Perhaps t h i s  constant  food  I  use. i s  value  and  availability. Studies  by R o c h e l l e  ( p e r s . comm.) show a v e r a g e c r u d e p r o t e i n  v a l u e s o f 20.5 and 9.9 pdm f o r d e e r f e r n f o r t h e May t o J u n e and July  to  September  Average crude f i b r e periods  of  respectively  ratios  a r e thus  f e r n f o r t h e above p e r i o d s ,  May  2.5 and  average crude p r o t e i n v a l u e s  to  Island.  ratios  June and J u l y t o September  bears  do n o t consume d e e r f e r n and sword  their  high  fibre:protein  ratios.  7.5).  same  Rochelle  o f 17.9 and 10.3 pdm f o r  and a v e r a g e  53.8 and 55.5 pdm f o r f i b r e : p r o t e i n  the  on V a n c o u v e r  l e v e l s were 50.4 and 74.7 pdm f o r t h e s e  (fibre:protein  a l s o found sword  periods  crude f i b r e  of 3.0 and 5.4 f o r  periods. fern  levels  may  The f a c t be  due  that to  91  The stalk in  l e a v e s o f d e v i l ' s c l u b were consumed by b e a r s , was n o t ( F i g . 2 3 ) .  t h e l e a v e s than  (Fig.  23).  (12-15)  than  in  animals  Total  are  highly  low  ash  determined  by  There of  nutrient  devil's  ( F i g . 25).  club  their  the  levels  consumed  by b e a r s  leaves  the  is  after  slightly  between t h e l e a v e s both  a p p e a r t o be  i s interesting  since  the  the choice o f the root  over  no  lower  throughout  apparent  tubers  and  ( F i g ^ 24).  three  the  differences leaves  of  green  (3) b e r r i e s .  of  higher  and  Crude  20 p e r c e n t early  (Fig.  19) .  coincidental  of and  huckleberry and i t s  protein  levels  of the dry matter  i n the s p r i n g .  Crude  ( F i g . 25 ) and a r e n o t  Crude p r o t e i n i n does  as  IV).  parts  (1) t h e newly g r o w i n g stem  time.  the  T h e r e a r e a l s o no  parts  s h a r p l y by J u n e  this  food  not decrease  the  other  t o t h e same  e x t e n t a s i n t h e n e w l y g r o w i n g stem d u r i n g June and A u g u s t . l e a v e s a r e consumed  and  o u t l i n e d f o r salmonberry  stem a r e a b o u t  decreased  of s o l u b l e  o f Mg, Ca, and K (App.  I sampled  d u r i n g May, and a r e p r o b a b l y protein  sampled  i n the stalk  levels  content  are  pattern  l e a v e s , and  i n t h e newly growing  The  composition  n u t r i e n t content;  (2) o t h e r  a l l seasons  Bears use t h i s  d i f f e r e n c e s i n the l e v e l s  follows  leaves,  in  proximate a n a l y s i s  in  higher  i n the stalk.  content  shifts  p r o t e i n was  was much h i g h e r  (1-3).  selective  huckleberry  for  ratio  and m o i s t u r e  volumes.  between n u t r i e n t  The  during  u s e o f skunk cabbage by b e a r s  at  apparent  of crude  f a t do n o t d i f f e r  parts of the plant.  season  stalk  leaves  i n the l e a v e s than  The  other  the  and c r u d e  stalk.  higher  the  The f i b r e : p r o t e i n  carbohydrate the  in  The l e v e l  while the  to berries  during  the  The  summer  92  Green pdm)  huckleberries  and c r u d e f i b r e  (2.5  pdm),  percent) (Fig.  25).  (6-8 pdm) levels  the  crude  values  fibre  of crude p r o t e i n  levels  (35  (15  pdm),  (82 p e r c e n t )  for  a crude  protein  24.8 pdm f o r r i p e ,  tall  level  crude  berries  o f crude  protein  comparatively  carbohydrates Mealey  6.1  found  i n Yellowstone  pdm;  fibre  (ADF),  Hamer e t a l .  pdm and a f i b r e  blue huckleberries  high  (NFE) (65  (1975)  a s h , 2.8 pdm). o f 6.1  fat  mature  scopariurn  9.7 pdm; e t h e r e x t r a c t ,  pdm; NFE, 67.8 pdm; and t o t a l  (1977) f o u n d  but  ( F i g . 25) .  Vaccinium  of  (19  pdm), and m o i s t u r e (82  values  f a t (11 pdm), s o l u b l e  f o r ripe  (crude p r o t e i n ,  of  carbohydrates corresponding  and c r u d e  pdm) , and m o i s t u r e  13.9  levels  R i p e h u c k l e b e r r i e s have low l e v e l s  of  similar  high  (35 pdm) , and l o w e r  soluble  than  had  i n Banff  level  National  Park. During are  close  Hay t h e c r u d e to  progresses,  24  crude  pdm  protein  ( F i g . 26,  protein  increases  slightly  relatively  c o n s t a n t a t about  The IV,  nutrient  since  I  and c r u d e f i b r e App. IV) .  decreases  (Fig. 26).  the  s a m p l e d them a t o n l y one t i m e .  10.84 pdm t o t a l  season  and c r u d e  remain  i n Appendix  T h e mean  crude fibre,  fibre  the season.  v a l u e s f o r c o h o salmon a r e i n c l u d e d  pdm s o l u b l e c a r b o h y d r a t e s , 5.6 pdm c r u d e  o f sedge  carbohydrates  40 pdm t h r o u g h o u t  v a l u e s f o r two coho f e m a l e s were 73.2 pdm  energy,  As  slightly  Soluble  levels  nutrient  protein,  7.95  4.99 k c a l s  gross  a s h , and 70.3 p e r c e n t m o i s t u r e .  93  6.3.3. The from  Food  amount  zero  to  availability during  availability of 274  kg  data  from  (396.2 k g )  per  hectare  applies  foods  to the  eaten  (4.3  berries  kg).  sample s i z e s  are very  are  in  common  the  of  by  low.  The  understory  (Table  bears. area  was  the  stink  These f i g u r e s  per h a b i t a t type  The  order of  were  20  overall  the  consumed  Ahnuhati  and  bear  are only estimates  the  food  Table  most abundant  currant  varied  The  i s shown i n  plant foods of  1).  p e r i o d s i n F i g u r e s 19  i n the study  salmon, salmonberry while  r  abundant  D  food a v a i l a b l e  w h i c h e a c h f o o d was  abundance o f b e a r Apart  bear  1. food  least  s i n c e the by  bears  watershed  (Table  -  6.4.  Discussion  6.4.1.  Comparison  To compare information (1973),  I  Clark  comparison,  the have (1957),  the  of food  observed reworked and  individual  c l a s s e s as f o l l o w s :  h a b i t s with seasonal the data  Mealey  other  food  areas  use  of Poelker  (1975)  to  existing  and  Hartwell  ( F i g . 27).  f o o d s p e c i e s have been lumped  To into  aid 12  94  Figure  27.  "  Comparsion of c l a s s e s of bear foods i n the Ahnuhati with e x i s t i n g studies of food h a b i t s of bears i n c o a s t a l ecosystems. . 1 2  g r a s s e s , s e d g e s , and horsetai1 herbs and f e r n s •  3  green  4  fruits  5  fish  6  small  mammals  7  large  mammals  8  unidentified foods  9  insects  10  shrub  fungus  : 11  wood  12  nuts  fibre  material  and  trace  >100n LU  > 50 H LU  DC  1 2  1 2  8  June Mealey  July (1975)  -  -  August  l a k e economy  N = 105 g r i z z l y  bear  scats  >100~f LU >  <  50H  LU  1 2  6 7 April Mealey  1 2 -  (1975)  June  6 July  -  8 9  1  6 7 8  September  - v a l l e y p l a t e a u economy  October N = 340 g r i z z l y  bear  -  10  November  scats  >100-| LU >  50H  LU  1 2  r—rr-i  6 7  1 2  June Mealey  (1975)  8 July  - m o u n t a i n economy  -  12_  2  September  12 October  .N = 170 g r i z z l y  bear  scats  > 100-1 u >  P 50< UJ £C  1 2 3  6 April  8  JZL  11  - June  ZlZL  4 July  P o e l k e r and H a r t w e l l  1 2  8 9 10 -  September  (1973)  6 October  288  black  -  8  10  November  bear stomachs  >100i UJ  >  50-  < i—r  UJ  1 2 3 4  1 2 3 4 5  8 May -  June This  July  8  -  September  study  J.].§„.t?J^.?.L.^D^...3D..^Jj...bear„..§.?.S.t;.?..,  >100i UJ  >  50-  < UJ  12  4 5 July  Clark  (1957)  -  October 140 g r i z z l y  bear  scats  CD  > 100-1  >  50-  LU  1 2  7 8  1 2  TpTiT"-' 1 'june Hamer e t a l .  1 2  July  -  2  September  (1978)  8 May -  7  N = 105 g r i z z l y  1 2 June  L l o y d and F l e c k  October -  (1977)  November  scats  8 9  4 July  bear  7  -  August 130 b l a c k  and g r i z z l y  bear  scats  to  98  Class  Food  1 2 3 4 5 6 7 8 9 10 11 12  All  g r a s s e s , s e d g e s , and h o r s e t a i l h e r b s and f e r n s green shrub m a t e r i a l fruits fish s m a l l mammals l a r g e mammals u n i d e n t i f i e d and t r a c e f o o d s insects fungus wood f i b r e nuts  o f t h e f o o d s r e p o r t e d by T i s c h  (1957),  The while  (1961), Mealey  Hamer e t a l . (1977), and L l o y d  t h e above  Type  and F l e c k  (1975),  Clark  (1977) f i t i n t o  classes.  comparison the  o f f o o d s used among  relative  study  areas  shows  l e v e l s o f some f o o d s a r e s i m i l a r ,  which  availability  of food rather than t o v a r i a t i o n s i n the preference  bears  (Fig. 27).  Mealey's  (Fig.  fruit  shrubs  of  i n the  fashion  the  the  "lake  for  the  (19 57)  July-October  Ahnuhati  not  whereas  occur  " l a k e economy" economy"  higher  may  use  of  diet  of  (Mealey account fish  on diet  lower Kodiak  level  feeding  the duration  and t i m i n g  area  September-October)  in  classes  the  plant  1975).  The l a c k o f  in  compensatory  (50  a  percent  IV)  in  (30 p e r c e n t IV) . o f use  Island  ( F i g . 27).  the  the Ahnuhati  grizzlies  commonly  with t h e Ahnuhati  f o u n d a much  July-September  (late  variations  example, t h e data from  and b e r r i e s ,  do  Y e l l o w s t o n e a s compared Clark  to  (1975) " l a k e economy" do n o t use t h e s e f o o d  27) s i n c e  communities  largely  For  show h i g h use o f s h r u b s in  due  there are  differences  of  are  that  This  of t h e r u n of sockeye when compared  of  salmon  than I found  in  i n the  i s due p r o b a b l y t o salmon i n  Claris  w i t h my s t u d y  area  99  (August-October). time  S i n c e salmon a r e n o t a v a i l a b l e  as b e r r i e s a r e r i p e ,  energy  source  berries  on t h e K o d i a k  at  the  a r e p r o b a b l y a more  Island  than  they  same  important  are i n the Ahnuhati  w a t e r s h e d. P o e l k e r and H a r t w e l l insects the it  and f u n g u s  diet  sampled  they  bears.  and  any s i g n  o f bears  fungus,  although  they  hibernation, of g r i z z l y  food supply The  essence  annual  energy  suggested t h e den.  d i f f e r e n c e , or  Hartwell  i n scats.  (1973)  I  highly  d i d not  mushrooms o r b r a c k e t  were common i n t h e s t u d y  section, and  area.  I discuss the implications  seasonal  supply  bears, using a energy  physiological energy  of  simple  model  to  on e v i d e n c e  in  the annual  t o be between mid-summer  evaluate  that  bears  majority  of  d u r i n g a s h o r t p e r i o d when  food i s a v a i l a b l e .  assimilation  nutrients to the  a d a p t a t i o n s whereby t h e  i s assimilated  o f body  demand.  o f t h e argument i s b a s e d  abundant, h i g h q u a l i t y for  levels i n  foods are probably  f e e d i n g on e i t h e r  i n terms of d a i l y  have e v o l v e d their  these  of  Introduction  In t h e f o l l o w i n g  nutrition  since  P o e l k e r and  use  I m p l i c a t i o n s of h i b e r n a t i o n t o feeding ecology  6.4.2.1.  size,  that  may be u n d e r - r e p r e s e n t e d  observe  6.4.2.  significant  T h i s may be a r e a l  t o the fact  stomachs,  digestible  observed  which d i d n o t a p p e a r a t s i g n i f i c a n t  of Ahnuhati  may be r e l a t e d  (1973)  The  critical  period  bioenergetic period i s  and t h e  fall  immergence  to  100  In  the  evolutionary  history  which c o u l d t a k e a d v a n t a g e quality to  f o o d would  of  a  of  seasonal  suggest  weight  that  generally in  such  changes,  hyperphagia.  seasonal  the period  i n late  the energy  sources a v a i l a b l e energy  July.  defecation  Roth  seven  scats  (sensu  day  apparently (Craighead  black  bears  reached  their  spite  of  in  their  T h i s suggests not  was  that  sufficient  to  (1975) n o t e d  Yukon d i d n o t add found  clear  day  from  fat  that  rapidly  until  seasonal differences i n i n S w i t z e r l a n d , w i t h low  November  August.  grizzly  through  June  Nelson  (1973,  and  peak o f  1978)  noted  b l a c k b e a r s and a H i m a l a y a n  black  summer.  and g r i z z l y  Folk  in  were  Pearson  among two A m e r i c a n  i n late Black  July  summer  i n t h e r e m a i n i n g months, w i t h a d i s t i n c t  per  hyperphagia bear  g a i n among  r a t e s o f c a p t i v e brown b e a r s  values  late  seasonal  and t h u s , a n i m a l s had t o draw on body  (1977)  v a l u e s o f two s c a t s p e r higher  that  seasonal  and  black bears u s u a l l y  i n spring  i n order t o survive.  bears i n the southwestern after  rates,  o f f o r b s and g r a s s e s .  balance  high  a n d come  place are  (1971) f o u n d  June o r e a r l y  consumption  reserves  defecation  that  extensive  maintain  of  The t h r e e a r e a s o f e v i d e n c e  of g r e a t e s t weight  They f o u n d  lowest weights  abundance  a d a p t a t i o n has t a k e n  J o n k e l and Cowan  Montana.  individuals  have i n c r e a s e d s u r v i v a l a n d f i t n e s s  d o m i n a t e t h e p o p u l a t i o n numbers.  which  the Ursidae,  1978)  bears i n North for  part  v a r y i n g with l a t i t u d e and C r a i g h e a d  able  to survive  and  by  of  America  and  annual  1972, L e n t f e r  f a t reserves  hibernate  the year, the l e n g t h of time  (Folk  snow  accumulation  e t a l . 1972).  h i b e r n a t i o n by l o w e r i n g t h e i r  mobilizing  typically  Bears a r e  metabolic  demand  1967, F o l k e t a l . 1972,  101  Nelson  1 973),.  season  o f the year and t h e i r  above  These f a t r e s e r v e s a r e p r o d u c e d  normal  during the a c t i v e  production reguires  requirements  for  growth,  energy  intake  maintenance,  and  reproduction. In  g e n e r a l , energy  extract  and  mid-summer density  less  fibre  content  Just  et  higher  a  relationship  dry  Mealey  basis  content, (1975)  food sources  Bears during  not  to  of  Since  fall  The Model  The  assumption  a  bear  crude  lower  caloric  t h e autumn  crude foods.  (1976) f o u n d  fibre  intake  an and  more f o o d on  s p r i n g , and b e c a u s e o f t h e h i g h e r wet  weiqht  would  be  suggested  are o f higher  bulky. that the  quality  than  summer. estimate  the  food  biomass  supply  begin  of  o b v i o u s l y exceeds a l l  abundance.  i s that the basal metabolic  does n o t d e p a r t  salmon, the  i n A u g u s t , t h e mid-summer  one o f h i g h e r f o o d  6. 4. 2. 2.  first  than  to  summer t h a n i n  would t h e r e f o r e need  t h e salmon r u n s  period i s also  a  difficult  (App. IV) and t h e i r  Hamer e t a l . (1977) a l s o  and e a r l y did  have  digestible  o f mid-summer and f a l l  we  more  W h i t t e m o r e and E l s l e y  abundance and d e n s i t y o f t h i s others.  content  the equivalent  and  of s p r i n q  While  foods  between  i n swine.  weight  moisture  those  moisture  and  both  i n s p r i n g and e a r l y  makes them l e s s  digestibility  is  The s p r i n g  a l . (1976)  inverse  food  abundant  and f a l l .  and  from  significantly  from  rate  (BMR)  K l e i b e r ' s (1961)  formula: BMR  = 70 W (kg)  (4)  102  then  the  BMR  o f a 200  kg g r i z z l y  b e a r i s 3.7  megacalories  per  day. The times  second assumption its  BMR  in  i s that  order  an a c t i v e mammal r e q u i r e s  t o m a i n t a i n energy  balance  2.5  (Gessaman  1973) . Data  a r e not a v a i l a b l e  hibernating  bears  information  on  (Folk  on o x y g e n c o n s u m p t i o n , quotients  given  substitute  variable  for  information  on  respiratory  rates  bear i s l e s s  t h a n BMR.  the  metabolic  for  discussion first  provide  hibernation,  earlier for  and  body  this  purposes,  energy  reproduction?"  Spencer  and  (EPD)  for  establish by  period  Hebert  The  data  heart  of a  hibernating  period  EPD  = 2.5  BMR  + 0.25  bear  would  hibernation  metabolism  the  spring  on  for  and,  foods of  maintenance,  weight i t  gain  is  in coastal  cited  important  1 to  B.C.  March  p e r s . comm.), t h e n t h e e n e r g y day  and  whether i t i s t h e o r e t i c a l l y  o f December  60  a  on  percent of the c o s t s  o c c u r but  grizzlies  the  as  BMR.  "Could 25  the  available  during  requirements  does n o t  to  that  t o use  The  and  0.85  to offset  i n a d d i t i o n to the  day  level.  of  without  a l . (1973)  i s below b a s a l  q u e s t i o n I examine i s ,  120  et  the metabolism  approximates  If hibernation the  metabolic  of a bear  discussion  includes  Nelson  I have assumed  demand  level  i t i s not p o s s i b l e  by  suggest t h a t  metabolic  Dnfortunately,  temperature,  suggests that t h i s  possible.  day  the  sufficient  qrowth,  actual  1978).  respiratory  The  the  31  generally (Leigh-  required  of  May  and  June  BMR  (120  days) ]/60  can  per be  determined:  a 200  kg  thus  [0.85  require  10.8  megacalories  (5) per  day  10 3  during t h i s The  period.  mean  salmonberry, moisture  caloric and d e v i l ' s  content  digestibility  i s  200 kg b e a r  food  f o r each  24 h o u r  Stubbs  (1975) t h a t  be  a l s o assumed  for  17  percent  to rate  a bear  to offset  i s unaffected  The  domestic  approximately  during t h i s  grizzly period.  1976)  then  (wet) o f t h i s  period  meat  in  from by  polar  Hunt and  bulk.  He  was 15 t o 20 p e r c e n t  swine l i t e r a t u r e d o e s n o t  i t does appear  25 p e r c e n t  by u s i n g e n e r g y  EPD = 2.5 BMR kg  mean  measures.  from  of  the  i t s spring  must a s s i m i l a t e ,  can be  The e n e r g y  possible energetic  food. therefore, a  energetic requirements  between J u l y and O c t o b e r .  (EPD) f o r t h i s  24 kg  evidence  stomach c a p a c i t y  minimum o f 75 p e r c e n t o f t h e i r  200  and c i t e d  bears i n t h e Ahnuhati  90 day p e r i o d  the  approximate  Elsley  f o r seal  these e s t i m a t i o n s are v a l i d , then  Grizzly  A  and  shoots of  and  (App. I V ) . I f we  approximately  o f passage  capacity  c o s t s of h i b e r n a t i o n  day  kcals/gm,  (Whittemore  24 h o u r s  weight.  stomach  sedge,  period.  t h a t a bear's  i t s body  If  ladyfern,  (1977) f o u n d r a t e s o f p a s s a g e  to  include  86  for  i s 4.56  would r e q u i r e  bears  of  club  a t 70 p e r c e n t  the  Best  density  during the  r e q u i r e d per  determined:  + 0.75 [ 0 . 8 5 BMR  would t h u s r e q u i r e  (120 days) ]/90 12.5 m e g a c a l o r i e s  (6) p e r day  104  6.4.2.3.  I m p l i c a t i o n s o f t h e model  Consider energy  how a b e a r  might  requirement  manage  from  the  Acknowledging the foods a v a i l a b l e period  ( F i g . 20),  and  using  of  t h e amount of f o o d  Equation The  diet, (39.9 (8.4  calculated  p e r c e n t ) , sedge percent),  digestibility (Hamilton these  fat  the  October  among  swine, one  can  which  October  make up t h e J u l y  skunk  et  leaving  (App. I V ) .  to  o f t h i s c h a p t e r , a r e salmon shoots  of  salmonberry  (8.3 p e r c e n t ) , l a d y f e r n and s p i n y wood  Berns  suggests  To e s t i m a t e reduced  to  demands  (31.4 p e r c e n t ) ,  and  Cursory  foods  berries  July  available.  r e q u i r e d t o meet t h e e n e r g y  the data  These r e l a t i v e  bears  berries.  the  observed  cabbage  salmon  with  examination  of  only the r e l a t i v e  v a l u e of  value  interest  and o t h e r s have abundant  the  t h a t salmon s h o u l d this  Since berries  v a l u e s a r e o f some  areas  the r e l a t i v e  percent).  fern  and  a l . (1977),  In f a c t ,  (3.0 for  1978) I c a n c o n s i d e r f u l l y  (1957),  grizzly  sources  daily  conversion e f f i c i e n c i e s f o r  i s known c u r r e n t l y o n l y  foods.  Clark  these  from  percent), berries  (5.5  food  this  6.  p r o p o r t i o n s of foods as  assimilate  during  v a r i o u s n u t r i e n t s which have been estimate  to  salmon  available  since  observed to  seek  nutrients i n  be o f g r e a t e r v a l u e  i s not the case. o f salmon and b e r r i e s ,  I have  t h e n u t r i e n t s w h i c h I measured i n t h e f o o d a c c o r d i n g  steps  of  The  relative  28  are  assimilation  efficiencies  those  Since both  the  f o r • the  s w i n e and  process  of F i n g e r l i n g  are  to  as shown i n F i g u r e 28.  (1914)  shown  conversion of n u t r i e n t s into  bears  than  omnivorous  in  Figure  body f a t .  monogastrics,  the  Quantity of nutrients  energy  i n feces  observed  in  food  apparent d i g e s t i b i l i t y of gross energy ( h u c k l e b e r r i e s - 8 8 . 4 7 ; salmon - 8 7 . 5 7 ( H a m i l t o n 1978))  energy l o s t to u r i n e (0.9kcal/g protein catab o l i z e d ( K l e i b e r 1961))  e n e r g y l o s t i n methane p r o d u c t i o n ( f o r swine 1 . 0 2 k c a l / g CHO ( B r e i r e m 1939))  heat o f  ure 28:  digestion  m e t a b o l i z a b l e energy E f f i c i e n c i e s f o r swine ( F i n g e r l i 1914) fat 93 p e r c e n t starch 83 p e r c e n t protein 74 p e r c e n t sucrose 72 p e r c e n t  net  Reduction of nutrients a v a i l a b l e energy f o r f a t p r o d u c t i o n .  energy  i n food  into  the net  106  corresponding markedly  efficiencies  from  The  these  nutrient  consisted  of  that  the  pdm.  On  reducing this  value into The 73.2 pdm  analysis  and  10.8  nutrient  pdm  crude  value  crude  levels  7.95  of  the  preceding  the  food are  salmon i s w o r t h body f a t . The greater  fat.  depart  huckleberries  65.1  Barker  pdm et  soluble  al.  the  soluble  found  pdm  7.6  carbohydrate  and  7.6  of  two  female  pdm  s o l u b l e carbohydrates,  to  scheme an  worth  2.8  sugar. coho salmon  kcal  averaged and  than  costs  berries  as  well?  its  5.6  to  salmon  carbohydrate  streams content  " q u i c k " energy  related  e t a l . 1976,  travelling  forage  of b e r r i e s  net  Elsley  enhance the  1976,  digestion  a  bears Lentfer  Elsley  away high  source  salmon  of  t o body  s w i n e i s known  of p r o t e i n t o e n e r g y  which do n o t o c c u r i n salmon.  do  Perhaps the  of p r o t e i n from  of s a l m o n  obviously  (1972) and  provide  of  V.  then  for berries.  Whittemore and  and  production  are  Why  o f body f a t i n d o m e s t i c  t o the i n t a k e r a t i o  energy,  several kilometers  ( F i g . 25)  f o r the conversion  accumulation  berries  Hensel  nutrient  a r e w o r t h 2.5,  f o r the  salmon.  B e r n s and  from  minerals  matter  handling  the c o s t s of handling  bears  also  of  salmonberries  g dry  of  ( p e r s . comm.) o b s e r v e d  The  r e d u c t i o n of t h e  estimate  2.8, per  of  A worked example i s d i s p l a y e d i n A p p e n d i x  energetic  consume  may  not  i n lowbush b l u e b e r r y averaged  carbohydrate analysis  mature  protein,  I have p a r t i t i o n e d  protein,  huckleberries  be  do  crude f a t . Using  fat.  probably  that  crude  pdm  sugar  pdm  showed  pdm  basis,  57.5  bears  values.  6.7  carbohydrates,  of  to  (Ehrensvard  1976).  Berries  or p r o v i d e v i t a m i n s  and  107  We need  can  now  to  October  salmonberry,  sedge,  on a v e r a g e ,  contribute  remaining  30  15  megacalories  period. fern,  percent  V) ,  and  25  v a l u e o f 577 k c a l demand  which  per day.  berries  would  from  require  Such i n t a k e  which,  i n 9 hours  for  of  required  848  during  the  the  shoots  July-October requirement  assuming  kcal  per  berries,  which  that  then the  to  period) o f 12.5 of  the  from  salmon,  wet  weight  have an  average  kg  meet  energy  its  daily  required f o r  f o r storaqe f o r hibernation, 9.2  to  of  (which t o g e t h e r make  energy  both  would  a  kq o f salmon and 4.5 kg o f  would r e p r e s e n t 5 p i n k salmon  according  our o b s e r v a t i o n s , c o u l d  and be  of f o r a g i n g .  Thus, i t does a p p e a r  possible  f o r a bear t o g a i n t h e  hibernation during the period  broader i m p l i c a t i o n s  of bears  the  further,  includes  grizzly  The  for  p e r kg wet w e i g h t ,  200  required  IV  And  percent  and t h e e n e r q y  collected  that  and skunk c a b b a g e  value  activity  berries  demand  10.6 m e g a c a l o r i e s , 75 p e r c e n t i s d e r i v e d  (App.  kq  energy  Assuming  of the daily  (mgakal).  h a s an a v e r a g e  energy  much o f t h e s e f o o d s a b e a r  i t s daily  percent  which  9400  how  t o consume t o s a t i s f y  July  up,  consider  of t h i s  of J u l y t o October.  to the ecology  are d i s c u s s e d i n the f o l l o w i n g  weight  chapter.  and  management  108  CHAPTER 7 - ADAPTATION OF BEARS TO ENERGY  Bears basic  are  a  successful,  O r s i d theme i s e x p r e s s e d  apparently  represent  environments. ruminants 1957,  in  and t h e y  various  exploit  be  Craighead  a high  Mealey  moisture  when  seasonal  v a r i a b l e environments  advantage  of  through  hyperphagic  subsequent The  the  restricted energy  will  capabilities  related  that  (Meyer-Holzapfel  highly  large  vulnerable  1975), and c a n o n l y  i t  low  fibre  and a h i g h d i g e s t i b i l i t y  (this  has  occurrence. only  a  These f o o d s o u r c e s Bears  because  availability  closely  individuals  of  they  of  (Nelson  survive i n can  take  abundant  1973,  tend  food  1978)  and  1967).  ( J o n k e l and Cowan 1971, R o q e r s  energy  different  r e p r o d u c t i v e performance of bears, i n p a r t i c u l a r  b e a r s , h a s been  follows  are  when  intake  h i b e r n a t i o n (Folk  which  They c a n e x p l o i t  Mysterud  i n their  seasonally  to  predators  1975, Hamer e t a l . 1 9 7 7 ) .  highly  strategies  digestive  they  1972,  content,  and  1978).  plant material e f f e c t i v e l y  content, study,  and  times  f a m i l y i n which t h e  responses  are not e f f e c t i v e  C u r r y - L i n d a h l 1972, H e r r e r o  (Craighead  forms  They do n o t have t h e  mammals a t o n l y t h o s e  to  cosmopolitan  AVAILABILITY  to their  nutritional  1977, B u n n e l l and T a i t  which a r e b e s t a d a p t e d  and produce f a t r e s e r v e s e s s e n t i a l  have t h e h i g h e s t f i t n e s s  (sensu Orians  for  condition 1 978).  It  to assimilate  winter  1969).  female  survival  109  This  kind  of  characteristics terms  of  tool,  but  of the l i f e  their  this  markinq  Bears during of  chapter,  I show how  which  i n the Ahnuhati  fail  (Nelson  to  shows  possess  adequate  fat  of  the  assimilation selected The  topography  interactions topography  and  space  6, t h i s  period  assimilation.  f a t reserves probably  ground  reserves  Bears will die  (Rogers  sguirrels  which  1976). do  die during hibernation and  lifestyle during  Morrison  critical  not  (Morton  1976).  which i n t e r f e r e s  the  of the Ahnuhati  among  bears  which  would  with  period  r e s u l t s i n a high  and,  approach is  be  i n f l u e n c e s movement  c o n t a c t among b e a r s  Ahnuhati  their  Any their  would  be  against.  encounter  valley  time  i n Chapter  during  p a r t i c u l a r , t h e r e e x i s t s the p o t e n t i a l  of  of bears,  r e p r o d u c t i v e output  that  bears'  of energy  treat  of f o o d .  in  i n terms o f energy  A r m i t a g e e t a l . 1976, G a l s t e r  aspects  of  As s u g g e s t e d  1973) o r h a v e r e d u c e d evidence  1975,  and abundance  sufficient  Indirect  to  m i q r a t i o n c o u l d be i n f l u e n c e d by  are concentrated  run.  gain  in  conceptual  data  the d i s t r i b u t i o n  their  the year i s c r i t i c a l  of  the  1971, Pyke e t a l . 1977).  and  salmon  amount  of  Ursidae  This process i s a useful  and t h e d i s t r i b u t i o n  the  evaluation  strategy of the  enormous  (Schoener  behavior,  topoqraphy  an  permits  history  fitness.  reguires  quantitatively In  deduction  vertical  in  interrupted (Chapter  of  expensive of bears  by  areas numerous  2, F i g . 2 ) .  salmon  frequent  along  The  the p r o b a b i l i t y  (1) The  tributary  in  aqgressive  valley  and t h e r i d g e l i n e  Movement  run  energetically.  and t h u s  i n two m a j o r ways: many  the  probability  creeks  walls  along the of  the  t h e watershed i s  110  t h u s c o n f i n e d t o e l e v a t i o n s below watershed in  by  a limited  b e a r s from  channels  watershed  a t one  and t r a v e l a  means  serve  contact. . river,  As  communication Rogers  of these p o i n t s , bottom  which  previously  selected  the  i s possible  only  areas  such  to the  valley  sites.  e n t e r the bottom,  probability  Bear  the  as  I f t h e r e were  the  for.  reduce  marking  probability  currents,  of  noisy  constrain  to large, s o l i t a r y  of  trees  t h e dense v e g e t a t i o n and  wind  pathways a v a i l a b l e  to  must  would r e d u c e  noted,  entry  bears  feeding  which  variable  {1977, p.99)  (2)  reach feeding  descend  t o the  scentposts  its  and  meadows,  t h e n i t would be as  with  To  sedge  o f communication  interactions, could  and  along the  m;  s u r r o u n d i n g watersheds  number o f p l a c e s .  spawning  700  the  mammals.  As  noted,  "in carnivores, scent g e n e r a l l y i s b e l i e v e d t o be o f s p e c i a l i m p o r t a n c e i n the a v o i d a n c e o f e n c o u n t e r s that could be m u t u a l l y damaging ( L e y h a u s e n and W o l f f 1959, H o r n o c k e r 1969, E a t o n 1970, E i s e n b e r g and K l e i m a n 1972, Ewer 1968, Mykytowycz 1974, P e t e r s and Mech 1 975).." In a r e a s clumped, (Fig.  where s e a s o n a l  o r have a l e s s dense  (Craighead 1976).  reduce  the  and C r a i g h e a d  number 1972,  I t i s possible that  established  w i t h marking  Intervalley also  can  cannot  give a quantitative  salmon,  viewed  available  because  established and  degree  Stonorov  and  through of  are  more  Ahnuhati  individual  interactions,  Stokes  1972,  dominance h i e r a r c h i e s c o u l d  movement o r m i g r a t i o n ( s e n s u be  food  Rogers also  be  trees.  1972)  net energy  of  v e g e t a t i o n cover than the  13), dominance h i e r a r c h i e s ,  interaction  in  concentrations  i n terms  of i n d i v i d u a l  e s t i m a t e of the  in a diet  t h e r e a r e no  Berns  relative  w i t h salmon  d a t a on  and  fitness.  I  difference  vs. a d i e t  the r e l a t i v e  Hensel  ability  without of  the  111  bears on  to gather  searching  consumption stomach  better  time,  of  worthwhile  their  digestibility.  It is be  salmon because o f salmon,  the  of  (high  caloric  bears'  annual  "packaging" basis). and  t h e energy  I n terms of t h e  thus  fitness,  i t  which h a s a  that  the  selective  i s probably  large  salmon  well run.  (1978) a r e c o r r e c t i n  advantages  of  sexual  i n bears are:  to  widely  and t h u s  t o provide competitive advantage t o l a r g e r  animals  in  allow  the  animal  l a r g e r would  which  between  comparatively  would  at  between  ranges  Schoener of  discrete,  1971, P o e l k e r  individual  u s e can be r e l a t e d  (after  and  v a l l e y s and which breeding.  Females  activity t o food  and/or  particularly Hartwell  which  plane. areas  and  the  abundance and  thus  1971, Pyke e t a l . 1977).  black  became  have an e n h a n c e d r e p r o d u c t i v e  a higher n u t r i t i o n a l  of spatial  home  mates, and  between  successful  valleys  overlap  fitness  travel  mates.  travelled  be more  through  to  more p o t e n t i a l  d i s p u t e s over  males  ability,  would  required to capture  inference  migrated  w i t h salmon  without  (1978) and H e r r e r o  encounter  Cowan  system,  Tait  2)  item,  digestive  and  1)  pattern  and  a diet  migrating to a r i v e r  dimorphism  The  through  per food  low h a n d l i n g t i m e  budget,  Bunnell  data are l a c k i n g  handling time  passage  palatability,  on a wet w e i g h t  bioenergetic  areas,  item,  f o r f a t p r o d u c t i o n than  density  to  of  Specifically,  t o assume, however, t h a t a d i e t  because  then  foods.  per food  rate  comparatively  and  If  time  capacity,  reasonable  the  different  grizzly  among  1973,  bears  females  Pearson  In some are  ( J o n k e l and  1975,  Sogers  112  1976).  Bunnell  predict  and  that discrete  resources  are  resource  the  prediction  bears  Craighead  patchy  and  first  providing interfere do  not  extensively  spatial  use  available  time  food  (MacArthur  ranked  should  for this  by  grizzlies  in  a c t u a l s p a w n i n g s i t e s and lower the thus  use  11  km  patches Pianka  distribution  use  of t h e i r  an  above to  black  1977,  Beecham bears  be and  Meslow  (1978)  in  Idaho  one  a c c o r d i n g to t h e i r  net  1966,  the  i n the  field.  c h o i c e of the  animal  (e.g. p r e d a t i o n )  of  a l l  optimal choice. are a v a i l a b l e i n clumps  ( F i g . 13).  do  The  of  the  The  I  data run,  salmon  only at  the the  extensive overlap i n  response  food.  not  Although  throughout  i n the Ahnuhati  adaptive  1971)  p e r i o d o f t h e salmon  occur  most c r i t i c a l  Schoener  of that type.  the  between i n d i v i d u a l s  considered  and  distribution  Ahnuhati  these  of the r i v e r  of s p a c e be  the  of  on  i n an a r e a ,  measurements  analysis during  i n the  and  among b l a c k  be t h e f i r s t  salmon s p a w n i n g s t r e a m s would be used  ranges  Reynolds  patch  d i s p r o p o r t i o n a t e l y with  data  (1978) a p p e a r s  (Lindzey  t h a t other e c o l o g i c a l f a c t o r s  quantitative  While  where  production.  and  with the  patch  should occur  p a t t e r n of a b e a r  compare t h i s  have  required  the  Tait  home  B e r n s e t a l . 1978) .  a l l  energy/feeding  The  areas,  ( p e r s . comm.)  are incomplete  and  unpredictable forage  order  then  of B u n n e l l  other  Bunnell  homogenous.  the o v e r l a p they observed  To p r e d i c t  and  and  distribution  overlap  1976,  attributed  could  In  and  (or t e r r i t o r i e s )  predictable  generally true. grizzly  (1978)  ranges  abundance and  studies,  to  Tait  to  river the  can  patchy  113  CHAPTER 8 - MANAGEMENT  Bears s u r v i v e adaptation section,  of  as  the stored demands  Throughout  their  energy on  for  As  needed  the  and  of  their  high  in  part  discussed  in  due the  f o r hibernation  animal  e v o l u t i o n , bears  concentrations  selected  omnivores,  hibernation.  energetic  locate  large  RECOMMENDATIONS  during  to  the  preceding  places  large  the  a c t i v e season.  which were a b l e  to effectively  quality  numbers  came  food to  were  probably  dominate  t h e bear  p o p u l a t i on. In r e l a t i v e l y have  been  added  recent as  environment o f bears. components quality crop  source  While dumps  n a t u r a l food  foods a r e s c a r c e . sources,  time, of  the  o r be e s p e c i a l l y  l o c a t e these  them.  a  garbage  may r i v a l  failure,  natural to  of  geologic  has  not  sources,  been  a p p e a r t o be c o n s i s t e n t w i t h  quality  i n the of  the  i n years o f  seasons  do b e a r s p o s s e s s  are a l s o capable  the feeding  man  quantified, this  during  o f b e a r s which o c c u r  of  food  particularly  important  but they  dumps  concentrated  nutritional  Not o n l y  The c o n c e n t r a t i o n s  garbage  when  adaptations  of r e l o c a t i n g  a t garbage  ecology  of  the  dumps family  Orsidae. It limited capable  is  predictable  success of  success  of  apparently  r e l o c a t i o n of bears  as a management t o o l  returning  rugged t e r r a i n  that  large  i n the world  deterrents  adverse  since  distances  (Hebert  bears  over  to  unpubl. d a t a ) .  obtain  have  are  only  clearly  some o f t h e most  also i s predictable since  conditions  will  natural  The  limited  bears t o l e r a t e foods.  This  114  suggests man  has I  of  a  high  not  m o t i v a t i o n t o o b t a i n f o o d , a m o t i v a t i o n which  been a b l e t o remove w i t h a wide r a n g e  suggest  that  t h e most e f f e c t i v e  g a r b a g e dumps, and  treat  the  treating  garbage  the  (e.g. b u r n i n g  bears  have n o t  habitat  protection.  Ahnuhati  f o r salmon runs  as r e s i d e n t therefore, and  and  bear  problems,  burial),  may  of  spawning  been i d e n t i f i e d , The  i s to  rather  annual  valleys,  than  probably i n  salmon.  The  migration  of  bears  that  coastal  feed bears  from  s u r r o u n d i n g watersheds  bears.  Watersheds l i k e  special  consideration  exact  b u t c o u l d be i m p o r t a n t f o r  suggests  grizzly  the  watersheds  Ahnuhati  to  with  may  the  large as  well  warrant,  f o r management o f b o t h  i s a s p e c i e s which i s s e n s i t i v e  of the n a t u r a l ecosystem Cowan  1972,  Herrero  resource  indefinitely,  (see f o r example  1976,  J o n k e l 1978).  development then  in coastal  the g r i z z l y  a n a t i v e s p e c i e s except  the  land  bear  i n park  S t o r e r and  I f the present  British  Columbia  probably w i l l  areas.  The  The  Fish  grizzly  bears  should  be  and  is  grizzly to  continues survive bear  management r e c o m m e n d a t i o n s which have d e v e l o p e d study  i t  not  1955,  pattern  specific  o f my  i f  Tevis  special  the course  consideration  to disturbance  receive  1)  use  people. The  of  bear-human  m i g r a t e between c o a s t a l  to the d i s t r i b u t i o n  routes used  salmon  the a s s o c i a t e d  of reducing  bears.  Grizzly response  way  of d e t e r r e n t s .  survive.  as  must Some during  are:  Wildlife  in coastal  Branch British  should determine Columbia  m a i n t a i n e d as h i g h d e n s i t y  and  grizzly  t h e demand f o r  d e c i d e which  areas  bear  These  areas.  115  areas  should  those  outlined  2)  consideration  significance  management  in  support  be  b)  have t h e i r  c)  be s u r v e y e d  large  the ecology  p e r s p e c t i v e , these  a)  runs  of g r i z z l y  rivers  and  past chronology  o f bear  o f salmon r u n s  use ( l i g h t ,  ground  by  the  Protection  actions  Regional  Biologist,  by  preceded  of the area  for  each  Wildlife and t h e i r  concerned..  area  Biologist, staff..  by  should  an The be  the Regional  Some s u g g e s t i o n s  management c o n s i d e r a t i o n s a r e :  identification migration  b)  compiled  medium, heavy)  reconnaissance  established  a)  From a  should  d u r i n g t h e s a l m o n r u n and c l a s s i f i e d  management  for special  bears.  identified  appropriate  Habitat  as  o f salmon have a  S p e c i a l management c o n s i d e r a t i o n s h o u l d be  aerial  such  i n 3) b e l o w .  degree  3)  management  C o a s t a l w a t e r s h e d s which  special bear  receive special  and  protection  of  inter-valley  routes  m i n i m i z a t i o n o f human-caused d i s t u r b a n c e d u r i n g t h e period  of  t h e salmon  bears  a t t h i s time  could  result  reserves  i s  run.  critical  i n a significant  with  Energy  conseguent  assimilation  and  curtail  during ii)  unnecessary  interference  r e d u c t i o n i n body f a t mortality  Managers m i g h t c o n s i d e r t h e f o l l o w i n g i)  by  vehicle  in  winter.  actions:  activity  near  streams  t h e salmon r u n  maintain  buffer  strips  of  mature  cover  along  116  streamsides.  The p o s s i b i l i t y  prone  s h o u l d be a c c o u n t e d  the iii)  areas  of  blowdown  in  wind  f o r i n the s i z e of  buffer strips,  post s i g n s  grizzly  which  bear  inform  activity  the  public  and e s t a b l i s h  of  nearby  rules of  conduct  i n the area. c)  maintain for  corridors  vertical  spawning strips the in  In  a)  by  these  t o the sides of the valley*  These  incinerators high  be  incinerators  15 m  grizzly  in  b)  In  should  access i s not possible. bear  areas,  be c o m p u l s o r y .  these  If their  cost  they  should  i s  being  s u b s i d i z e d by t h e g o v e r n m e n t . areas  established,  where  onset  of  pit  dumps  which  association  human  settlement  use o f i n c i n e r a t o r s  the  not  tall.  self-contained  t o s m a l l o p e r a t o r s i s p r o h i b i t i v e , then be  Timber  disposal.  burned  should  along  open p i t g a r b a g e dumps  t o which bear  density  kilometer  1 chain i n width.  Columbia,  should  every  be a t l e a s t  a s a means o f r e f u s e  Garbage  from  cover salmon  should  British  provide the  and be a t l e a s t strips  which  bears  should occur a t l e a s t  coastal  In  access  areas  river  abandoned  of timber  must c o i n c i d e w i t h  a l l garbage d i s p o s a l .  by  be p e r m i t t e d .  will the  result  bears  in  o f man w i t h  Interim an food  open  initial should  117  5)  Illegal  apparent stiffer of  self  bears  Service  killed  the  should  as  does  not  occur  for  be d e t e r r e d w i t h  much  which p r e s e n t l y e x i s t .  self  defence  All  parts  s h o u l d a u t o m a t i c a l l y become  should not occur  within  and W i l d l i f e  salmon  run,  of t r e e cover  overnight  estuaries,  meadows  o r such  spatial  management  should  where r o a d s  sedge  areas  2  km  of  salmon  B r a n c h and F o r e s t  permit i t .  camping  w i t h i n 2 km o f salmon spawning  A strip  The  those  unless the Fish  explicitly  screen i n areas  9)  which  property.  During  such  bears,  reasons,  for  channels  permitted  8)  defence  Human s e t t l e m e n t  spawning  7)  of grizzly  p e n a l t i e s than  government  6)  kill  be  maintained  those  n o t be  as  visual  channels.  are adjacent  like  should  a  t o known f e e d i n g  o f t h e Sim and  areas,  Klinaklini  as snowslides.  and  temporal  should  second growth f o r e s t  cut  pattern  recognize  i s probably  of  within  special  t h a t t h e 15 t o 40 y e a r o l d very  low  productivity  to  bears.  10)  In  s p e c i a l management a r e a s , c o n t r o l o f r o a d  the hunting Wildlife  season  Branch  s h o u l d be an o p t i o n and r o a d a l t e r a t i o n  be t h e r e s p o n s i b i l i t y  open  to  which l i m i t s  of the logging c o n t r a c t o r s  access the  during  Fish  access  and  should  concerned.  118  11)  F u r t h e r study  bears in  of the i n f l u e n c e of c l e a r c u t t i n g  i s essential.  Rivers Inlet  The work i n i t i a t e d  provides  work s h o u l d be s u p p o r t e d  an i d e a l  by B.C.  experimental  and c o n t i n u e d .  on  grizzly  Forest  Products  situation.  This  119  LITERATURE  CITED  Amstrup, S.C. and J . Beecham. 1976. A c t i v i t y p a t t e r n s of r a d i o - c o l l a r e d b l a c k b e a r s i n IdahoJ . W i l d l . Manage. 40(2) : 340-348. A r m i t a g e , K.B., J . F . Dowhower, and G.E. S v e n d s e n . 1976. S e a s o n a l c h a n g e s i n w e i g h t s o f marmots. Am. M i d i . Nat. 36-51.  96:  A s s o c i a t i o n o f O f f i c i a l A g r i c u l t u r e C h e m i s t s (AOAC). 1970. O f f i c i a l methods o f a n a l y s i s . 11th e d . , W a s h i n g t o n , D.C. Atwell,  G., D. Boone, J . G u s t a f s o n , and V.D. B e r n s . In p r e s s . Brown b e a r summer a l p i n e h a b i t a t r e q u i r e m e n t s on t h e Kodiak N a t i o n a l W i l d l i f e Refuqe i n P r o c e e d i n q s o f the 4th I n t e r n a t i o n a l C o n f . on B e a r R e s e a r c h and Management. C.J. M a r t i n k a , e d . , K a l i s p e l l , Montana 1977.  B a r k e r , W.G., F.A. Wood, and W.B. Collins. 1963. Sugar l e v e l s i n f r u i t s of t h e lowbush b l u e b e r r y e s t i m a t e d a t f o u r p h y s i o l o g i c a l a g e s . N a t u r e 198: 4882. Beeman, L . E . and M. P e l t o n . In p r e s s . S e a s o n a l f o o d s and f o o d e c o l o g y o f t h e b l a c k bear i n t h e Smoky M o u n t a i n s i n 4 t h I n t e r n a t i o n a l C o n f . on B e a r R e s e a r c h and Management. C.J. M a r t i n k a , ed. K a l i s p e l l , Montana. 1977,. B e r n s , V.D. and R . J . H e n s e l . 1972. R a d i o - t r a c k i n g brown b e a r s on K o d i a k I s l a n d . Pp. 19-25 i n S. H e r r e r o , ed. Bears t h e i r b i o l o g y and management. IOCN New S e r i e s No. 23, Morges, S w i t z e r l a n d . B e r n s , V.D., G.C. A t w e l l , and D.L. Boone. In p r e s s . F a l l brown b e a r h a b i t a t use and movements o f K a r l u k Lake on K o d i a k I s l a n d . I n C . J . M a r t i n k a , ed. B e a r s - t h e i r b i o l o g y and management. B e s t , R. 1977. E c o l o g i c a l a s p e c t s of p o l a r bear n u t r i t i o n . Pp. 203-211 i n R.L. P h i l i p s and C . J . J o n k e l , e d s . P r o c e e d i n g s o f the 1975 P r e d a t o r Symposium June 16-19, 197 5. Montana F o r e s t and C o n s e r v a t i o n E x p e r i m e n t S t a t i o n , Missoula. i x + 268 pp. B o u l i e r e , F. 1954. The n a t u r a l h i s t o r y t r a n s l a t i o n from French. A l f r e d A. 387 pp. B r e i r e m , K. 1939. Tierernahrung  Der 11:  Energieumsatz 487-528 c i t e d  o f mammals, 1970 K n o p f , New York.  xxi +  b e i den S c h w e i n e n . by K l e i b e r (1961).  B u n n e l l , F.L. and D.E.N. T a i t . 1978. P o p u l a t i o n dynamics of b e a r s and t h e i r i m p l i c a t i o n s . P r o c . o f I n t . C o n f . on Pop. Dynamics o f L a r g e Mammals, May 1978, L o g a n , U t a h .  120  B u r k h a r d t , D., W. S c h l e i d t , a n d H. A l t u e r . 1966. Signals i n the a n i m a l w o r l d , 1967 t r a n s l a t i o n o f S i g n a l e i n der Tierwelt. M c G r a w - H i l l , New Y o r k . 150 pp. C a h a l a n e , V. 1947. The b e a r s - b l a c k b e a r . Pp. 134-143 i n Mammals o f N o r t h A m e r i c a . The M a c M i l l a n Co., New Y o r k . 578 pp. C i t e d by S h a f f e r (1971). C h a t e l a i n , E.F. Peninsula.  1950. Bear-moose r e l a t i o n s h i p s on t h e K e n a i T r a n s . N. Am. W i l d l . C o n f . 15: 224-234.  C h e r r y , J . S . and M.E. P e l t o n . 1976. R e l a t i o n s h i p s between body measurements and w e i g h t o f t h e b l a c k b e a r . J . Tenn. A c a c . S c i . 51 ( 1 ) : 32-34. C l a r k , W.K. Trans. Cole,  1957. N. Am.  Seasonal f o o d h a b i t s of t h e Kodiak W i l d l . C o n f . 22: 145-151..  bear.  G.F. 1972. G r i z z l y b e a r - e l k r e l a t i o n s h i p s i n Y e l l o w s t o n e N a t i o n a l Park. J . W i l d l . Manage. 3 6 ( 2 ) : 556561.  Cowan, I . McT. 1938. G e o g r a p h i c d i s t r i b u t i o n o f c o l o r p h a s e s o f t h e r e d f o x and b l a c k bear i n t h e P a c i f i c N o r t h w e s t . J. Mammal. 19 ( 2 ) : 202-206. Cowan, I . McT. 1972. S t a t u s and c o n s e r v a t i o n o f b e a r s o f t h e world — 1970. Pp. .343-346 i n S. H e r r e r o , e d . B e a r s t h e i r b i o l o g y and management. IUCN New S e r i e s No. 23, Morges, S w i t z e r l a n d . C r a i g h e a d , F.C. J r . 1976. G r i z z l y b e a r r a n g e s and movement a s d e t e r m i n e d by r a d i o t r a c k i n g . Pp. 97^-109 i n M.R. P e l t o n , J.W. L e n t f e r , and G.E. F o l k , e d s . B e a r s - t h e i r b i o l o g y and management IUCN New S e r i e s No- 40, Morges, S w i t z e r l a n d . C r a i g h e a d , F . C , J r . and J . J . C r a i g h e a d . 1972.. Data on g r i z z l y b e a r d e n n i n g a c t i v i t i e s and b e h a v i o u r o b t a i n e d by u s i n g w i l d l i f e t e l e m e t r y . Pp. 84-106 i n S. H e r r e r o , e d . B e a r s t h e i r b i o l o g y and management IUCN New S e r i e s No. 23, Morges, S w i t z e r l a n d . C r a i g h e a d , J . J . , J.R. V a r n e y , and F.C. C r a i g h e a d , J r . 1974. A population a n a l y s i s o f the Yellowstone g r i z z l y bears. Montana F o r e s t and C o n s e r v a t i o n E x p e r i m e n t S t a t i o n , S c h o o l o f F o r e s t r y , U n i v e r s i t y o f Montana, M i s s o u l a , B u l l . No. 40. 20 pp. C u r r y - L i n d a h l , K. 1972. The brown b e a r ( U r s u s a r e t o s L.) i n E u r o p e : d e c l i n e , p r e s e n t d i s t r i b u t i o n , b i o l o g y , and ecology. Pp. 74-81 i n S. H e r r e r o , e d . B e a r s - t h e i r b i o l o g y and management. IUCN New S e r i e s No. 23, Morges, Switzerland. Darke, L . P e r s . comm. C a n a d i a n K n i g h t I n l e t , B.C.  Federal Fisheries  patrolman.  121  D a v i s , D.D. 1964. The g i a n t p a n d a , a m o r p h o l o g i c a l s t u d y o f e v o l u t i o n a r y mechanisms. F i e l d i a n a Z o o l . Mem., V o l . 3. C h i c a g o Nat. H i s t . Mus. 339 pp. Day,  B. 1957. G r i z z l i e s i n t h e i r b a c k y a r d . I n c . , New Y o r k . 224 pp.  E a t o n , R.L. 1970. territoriality  J u l i a n Messner  Group i n t e r a c t i o n s , s p a c i n g , i n c h e e t a h s . Z. T i e r p s y c h o l .  Edwards, R.Y. and D.E- G r e e n . t o census g r i z z l y bears.  and 27: 481-491.  1959. The measurement o f t r a c k s M u r r e l e t 4 0 ( 2 ) : 14-16.  E h r e n s v a r d , Y., F. B e r s c h a u e r , K.H. Menke, E. R o g d a k i s , and G. Sturm. 1976. E f f e c t o f r a t i o n c o m p o s i t i o n on p l a s m a i n the g r o w i n g p i g . Pp. 85-88 i n M. V e r m o r e l , e d . E n e r g y m e t a b o l i s m o f f a r m a n i m a l s . EAAP P u b l i c a t i o n No. 19. G. de B u s s a c , C l e r m o n t - F e r r a n d , x v i + 364 pp. E i s e n b e r g , J . F . 1966. The s o c i a l o r g a n i z a t i o n o f mammals. Handbuch d e r Z o o l o g i e , p a r t 10, 8 ( 2 6 ) : 1-92. B e r l i n : De Gruyter. E i s e n b e r g , J . F . and D.G. K l e i m a n . 1972. O l f a c t o r y c o m m u n i c a t i o n i n mammals. Ann. Rev. E c o l . S y s t .  3: 1-32.  E l s l e y , F.W.H. 1976. P r o t e i n n u t r i t i o n i n t h e b r e e d i n g sow. Pp. 353-367 i n C o l e , D.J. A., K.N. Boorman, P . J . B u t t e r y , D. L e w i s , R . J . N e a l e , a n d H. Swan. 1976. Protein m e t a b o l i s m and n u t r i t i o n . E u r o p e a n A s s o c i a t i o n f o r A n i m a l P r o d u c t i o n P u b l . No. 16. B u t t e r w o r t h s , London. 0 + 515 pp. Ewer, R.F. York.  1968. E t h o l o g y o f mammals. Plenum 418 pp. C i t e d by R o g e r s ( 1 9 7 7 ) .  Fingerling, . 1914. referenced.  Cited  by K l e i b e r  Press.  New  (1961) b u t n o t  F l e c k , S., K. L l o y d , and B. S m i t h . 1977. Some d i s t i n g u i s h i n g c h a r a c t e r i s t i c s of plant p a r t s found i n c o a s t a l bear d r o p p i n g s . B.C. F i s h and W i l d l i f e B r a n c h R e p t . , Nanaimo. 10 pp. F l o w e r s , R. 1977. The a r t and t e c h n i q u e o f s n a r i n g b e a r s . Washington F o r e s t P r o t e c t i o n A s s o c i a t i o n , S e a t t l e . 37 pp. Folk,  G.E. 1967. P h y s i o l o g i c a l a d a p t a t i o n s o f s u b a r c t i c b e a r s under w i n t e r den c o n d i t i o n s Pp. 78-85 i n K. F i s h e r and F. S o u t h , e d s . Mammalian H i b e r n a t i o n I I I : P r o c . o f t h e T h i r d I n t . Symp., S e p t . 13-16, 1975.  Folk,  G. E., J r . , M. F o l k , and J . J . M i n o r . 1972. Physiological c o n d i t i o n o f t h r e e s p e c i e s o f b e a r s i n w i n t e r dens. Pp. 107-124 i n S. H e r r e r o , e d . B e a r s - t h e i r b i o l o g y and management. IITCN New S e r i e s 23, Morges, Switzerland.  122  Frame, G.W. Creek, Gard,  1974. B l a c k bear p r e d a t i o n on s a l m o n a t O l s e n Alaska. Z. T i e r p s y c h o l . 35:23-28.  R. 1971; Brown b e a r p r e d a t i o n on s o c k e y e salmon a t K a r l u k L a k e , A l a s k a . J . W i l d l . M a n a g e . . 3 5 ( 2 ) : 193-204.  Gessaman, J.A. 1973. Methods o f e s t i m a t i n g e n e r g y c o s t o f f r e e existence. Pp. 3-31 i n J.A. Gessaman, e d . Ecological e n e r g e t i c s o f homeotherms: a view c o m p a t i b l e w i t h e c o l o g i c a l m o d e l i n g . . V o l . 20. Utah S t a t e U n i v . P r e s s , Logan. 155 pp. G l e n n , L.P. and L.H. M i l l e r . I n p r e s s . S e a s o n a l movements o f an A l a s k a n p e n i n s u l a brown b e a r p o p u l a t i o n . I n - C . J . M a r t i n k a , ed. B e a r s - t h e i r b i o l o g y and management. Haglund, and the  B. 1968. W i n t e r h a b i t s o f t h e b e a r ( U r s u s a r c t o s L.) t h e w o l f ( C a n i s l u p u s L.) a s r e v e a l e d by t r a c k i n g i n snow. V i l t r e v y 5 ( 6 f : 213-361.  Hamer, D., S. H e r r e r o , and R.T. O g i l v i e . 1977. Ecological s t u d i e s o f t h e Banff N a t i o n a l Park g r i z z l y bear: Cuthead Wigmore R e g i o n , 1976. P a r k s Canada C o n t r a c t WR-34-76. 239 PPHamer, J.D.W. 1974. D i s t r i b u t i o n , a b u n d a n c e , and management i m p l i c a t i o n s o f t h e g r i z z l y b e a r and mountain c a r i b o u i n t h e M o u n t a i n C r e e k w a t e r s h e d o f G l a c i e r N a t i o n a l P a r k , B.C. U n p u b l . M.Sc. T h e s i s , U n i v . o f C a l g a r y , C a l g a r y , A l b e r t a . 164 pp. Hanson, W.R. and F. G r a y b i l l . 1956. Sample s i z e analysis. J . W i l d l . Manage. 20: 64-68.  i n food-habits  H a t l e r , D.F. Alaska.  1972. Food h a b i t s o f b l a c k b e a r s i n i n t e r i o r Can. F i e l d Nat. 8 6 ( 2 ) : 17-31 I l l u s .  H e b e r t , D.M. Wildlife  P e r s . comm. R e g i o n a l w i l d l i f e B r a n c h , Nanaimo, B.C.  biologist.  Fish  and  H e d i g e r , H. 1946. Raum-zeit s y s t e m d e r t i e r e ( A r e a - t i m e s y s t e m o f a n i m a l s ) . S c h w e i z , Z. P s y c h o l . 5: 241. C i t e d by MeyerH o l z a p f e l (1957). H e n s e l , R . J . , W.A. T r o y e r , and A.W. E r i c k s o n . 1969. R e p r o d u c t i o n i n t h e f e m a l e brown b e a r . J . W i l d l . 3 3 ( 2 ) : 357-365. H e r r e r o , S. 1970.. Man and t h e g r i z z l y b e a r future?). B i o s c i e n c e 20 ( 1 ) : 1148-1153.  Manage.  ( p a s t , p r e s e n t , but  H e r r e r o , S. 1976. C o n f l i c t s between man and g r i z z l y b e a r s i n the N a t i o n a l Parks of North America. Pp. 121-145 i n M.. P e l t o n , J.W. L e n t f e r , and G.E. F o l k , e d s . B e a r s - t h e i r b i o l o g y and management.  123  H e r r e r o , S1978. A c o m p a r i s o n o f some f e a t u r e s o f t h e e v o l u t i o n , e c o l o g y , and b e h a v i o r o f b l a c k and g r i z z l y bears. C a r n i v o r e 1 ( 1 ) : 7-17. H o l z w o r t h , J . M. 1930. Putnam's Sons, New H o r n o c k e r , M.G. J. . W i l d l .  The w i l d g r i z z l i e s o f A l a s k a . York. x x i + 417 pp.  1969. Winter t e r r i t o r i a l i t y Manage. 3 3 : 457-464.  brown  G.P.  i n mountain  lions.  Hunt, J.N. and D.F. S t u b b s . 1975. The volume and e n e r g y c o n t e n t o f meat as d e t e r m i n a n t s o f g a s t r i c e m p t y i n g . P h y s i o l . (London) 245(1) : 209-225. J o n k e l , C.J. p e r s . comm. D e p t . Montana, M i s s o u l a 59801.  of Forestry,  University  J. of  J o n k e l , C.J. 1977. Annual r e p o r t - border g r i z z l y p r o j e c t , 1976. S c h o o l o f F o r e s t r y , U n i v e r s i t y o f Montana. No. 2. Jonkel, C.J. 1978. Annual Beport - border g r i z z l y p r o j e c t . S c h o o l o f F o r e s t r y , U n i v e r s i t y o f Montana. No. 3. 134 pp. J o n k e l , C . J . and I . McT. spruce-fir forest.  Cowan. 1971. The b l a c k W i l d l . Monog. 27: 1-57.  bear i n t h e  J o n k e l , C . J . and C.W. Servheen. 1977. B e a r s and p e o p l e : A wilderness challenge. W e s t e r n W i l d l a n d s . 4 ( 2 ) : 22-25. Just,  A., O.K. Easmussen, and H.L. Hansen. 1976. Factors i n f l u e n c i n g t h e d i g e s t i b i l i t y and e f f i c i e n c y o f u t i l i z a t i o n o f m e t a b o l i z a b l e e n e r g y (ME) i n d i e t s o f g r o w i n g p i g s . Pp. 289-292 i n M. V e r m o r e l , e d . EAAP Pub. No. 19.  K l e i b e r , M. York.  1961. The f i r e x x i i + 454 pp.  K l e i n , D.E. 1959. populations.  of l i f e .  John  W i l e y and  Sons,  New  Track d i f f e r e n t i a t i o n f o r c e n s u s i n g bear J . W i l d l . Manage. 23 ( 3 ) : 36 1-363.  K i s t c h i n s k i i , A.A. 1972. L i f e h i s t o r y o f t h e brown b e a r ( U r s u s a r c t o s L.) i n n o r t h e a s t e r n S i b e r i a . Pp..67-73 i n S. H e r r e r o , ed. B e a r s - t h e i r b i o l o g y and management. IUCN New S e r i e s P u b l . 23. Krajina, V.J. 1965. The b i o g e o c l i m a t i c c l a s s i f i c a t i o n of B r i t i s h Columbia. A m e r i c a 1: 1-17.  z o n e s and E c o l . Western  K r o t t , P. and G. K r o t t . 1963. Zun v e r h a l t e n ( U r s u s a r c t o s L. 1758) i n den A l p e n . . Z. 160-206. C i t e d by E i s e n b e r g ( 1 9 6 6 ) .  N.  des b r a u n b a r e n T i e r p s y c h o l . 20:  K u r t e n , B. 1976. The c a v e b e a r s t o r y . L i f e and d e a t h o f a v a n i s h e d a n i m a l . C o l u m b i a U n i v . P r e s s , New Y o r k . 163 pp.  124  L a n g i n , H. p e r s . comm. Wildlife W i l d l i f e B r a n c h , Nanaimo.  T e c h n i c i a n , B.C. F i s h and  L e i g h - S p e n c e r , S. p e r s . comm. Biologist, Box 130, C r o f t o n , B.C. VOE 1R0. L e n t f e r , J . p e r s - comm. Biologist. Game, J u n e a u , A l a s k a . 99801.  B.C. F o r e s t  Alaska  Products,  D e p t . o f F i s h and  L e n t f e r , J.W., R . J . H e n s e l , L.H. M i l l e r , L.P. G l e n n , and V.D. Berns. 1972. Remarks on d e n n i n g h a b i t s o f A l a s k a brown bears. Pp. 125-132 i n S. H e r r e r o , e d . B e a r s - t h e i r b i o l o g y and management. L e y h a u s e n , P. a n d R. W o l f f . 1959. Das r e v i e r e i n e r h a u s k a t z e . Z . . T i e r p s y c h o l 16: 666-670. C i t e d by R o g e r s (1977). L i n d e m a n n , W. 1954. M i t t . (Mammalogy  B r a u n b a r e n (Brown b e a r s ) . B u l l e t i n ) 2,1.  L i n d z e y , F.G. and E.C. Meslow. o f b l a c k b e r s on an i s l a n d 41 (3) : 408-412.  Saugetier  Kundl  1977. P o p u l a t i o n c h a r a c t e r i s t i c s i n W a s h i n g t o n . J . W i l d l . Manage.  L l o y d , K.A- a n d S. F l e c k . 1977. Some a s p e c t s o f t h e e c o l o g y o f b l a c k a n d g r i z z l y b e a r s i n s o u t h e a s t e r n B.C. B.C. F i s h and W i l d l i f e B r a n c h R e p o r t , C r a n b r o o k , B.C. 55 pp. M a c A r t h u r , R.H. and E.R. P i a n k a . 1966. On o p t i m a l u s e o f a patchy environment. Am. Nat. 100: 603-609.. M a r t i n k a , C . J . P e r s . comm. Research N a t i o n a l P a r k , Montana.  biologist.  Glacier  M a r t i n k a , C . J . 1971. S t a t u s and management o f g r i z z l y b e a r s i n G l a c i e r N a t i o n a l P a r k , Montana. T r a n s . N. Am. W i l d l . N a t . R e s o u r c e C o n f . 36: 312-322. M a r t i n k a , C . J . 1972. H a b i t a t r e l a t i o n s h i p s o f g r i z z l y b e a r s i n G l a c i e r N a t i o n a l P a r k , Montana. N a t i o n a l Park Serv. Unpubl. manuscript. 19 pp. M a r t i n a , C . J . 1976. E c o l o g i c a l r o l e and management o f g r i z z l y b e a r s i n G l a c i e r N a t i o n a l P a r k , Montana. Pp. 147-156 i n M.R. P e l t o n , J.W. L e n t f e r , a n d G.E. F o l k , e d s . B e a r s t h e i r b i o l o g y and management. M e a l e y , S.P. 1975. The n a t u r a l f o o d h a b i t s o f f r e e r a n g i n g g r i z z l y b e a r s i n Y e l l o w s t o n e N a t i o n a l P a r k , 1973-1974. U n p u b l . . M. Sc. T h e s i s , Montana S t a t e U n i v e r s i t y , Bozeman, Montana. 158 pp. Meehan, W.R. 1961. O b s e r v a t i o n s behaviour of g r i z z l y bears.  on f e e d i n g h a b i t s and Am. M i d i . N a t . 6 5 ( 2 ) :  409-412.  125  Meyer-Holzapfel, M. 1957. Handbuch d e r Z o o l o g i e  Dem v e r h a l t e n d e r Baren V I I I 1 0 ( 1 7 ) : 1-28.  M i l l s , E.A. 1919.. The g r i z z l y . The R i v e r s i d e C a m b r i d g e , New Y o r k . 289 pp.  (Ursidae) .  Press,  M o r r i s o n , P. and W. G a l s t e r . 1975. P a t t e r n s of h i b e r n a t i o n i n t h e A r c t i c g r o u n d s g u i r r e l . Can. J . Z o o l . 53: 1345-1355. M o r t o n , M.L. 1975. S e a s o n a l c y c l e s o f body w e i g h t s and l i p i d s i n B e l d i n g g r o u n d s q u i r r e l s . B u l l . S. C a l i f . Acad. S c i . 74: 128-143. M u e l l e r - D o m b o i s , D. and H. E l l e n b e r g . 1974. Aims and methods of vegetation ecology. John W i l e y and S o n s , New Y o r k . xx + 547 pp. M u l l e r - S c h w a r z e , D; 1974. S o c i a l f u n c t i o n s o f v a r i o u s s c e n t g l a n d s i n c e r t a i n u n g u l a t e s and t h e p r o b l e m s e n c o u n t e r e d i n e x p e r i m e n t a l s t u d i e s o f s c e n t c o m m u n i c a t i o n . Pp. 107-114 i n V o l . 1. 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. IUCN P u b l . New S e r i e s 24. Morges, S w i t z e r l a n d . Mundy, K.R.D. 1963. E c o l o g y o f t h e g r i z z l y b e a r ( U r s u s a r c t p s L.) i n G l a c i e r N a t i o n a l P a r k , B r i t i s h C o l u m b i a . M.Sc. T h e s i s , U n i v . o f A l b e r t a , Edmonton. 103 pp. M u r i e , A. 1944. The w o l v e s o f Mount M c K i n l e y . Fauna o f t h e n a t i o n a l parks o f the United States. Fauna S e r i e s No. 5, U.S. G o v t . P r i n t i n g O f f i c e , W a s h i n g t o n , D.C. 238 pp. M u r i e , 0. 1954. A f i e l d M i f f l i n Co., B o s t o n -  guide t o animal t r a c k s . 374 pp.  Houghton  Mykytowycz, R. 1974. Odor i n t h e s p a c i n g b e h a v i o r o f mammals. Pp. 327-343 i n M.C. B i r c h , e d . Pheremones. American E l s e v i e r P u b l i s h i n g Co., I n c . New York. 495 pp. Mysterud,, I . 1975. Sheep k i l l i n g and f e e d i n g b e h a v i o r o f t h e brown b e a r ( U r s u s a r c t p s ) i n T r y s i l , s o u t h Norway 1973. Norw. J . Z o o l . 23: 243-260. Nagy, J.A. and R.H. R u s s e l l . 1978. E c o l o g i c a l s t u d i e s o f t h e b o r e a l f o r e s t g r i z z l y bear (Ursus a r c t p s L . ) . Annual r e p o r t f o r 1977. C a n a d i a n W i l d l i f e S e r v i c e . 72 pp. N e l s o n , R.A. 1973. W i n t e r s l e e p P r o c . 48: 433:437.  i n t h e black  bear.  Mayo  Clin.  N e l s o n , R.A. 1978. Urea m e t a b o l i s m i n t h e h i b e r n a t i n g b l a c k b e a r . K i d n e y I n t e r n a t i o n a l 13 Supplement 8: 177-179. N e l s o n , R.A., H.W. W a l i v e r , J.D. J o n e s , R.D. E l l e f s o n , and P.E. Zollman. 1973. M e t a b o l i s m o f b e a r s b e f o r e , d u r i n g , and a f t e r winter sleep. Am. J . P h y s . 224 ( 2 ) : 491-496.  126  O r i a n s , G.H. 1969. On t h e e v o l u t i o n o f m a t i n g and mammals. Am. N a t . 103: 589-603.  systems  i n birds  P e a r s o n , A.M.. 1975.. T h e n o r t h e r n i n t e r i o r g r i z z l y b e a r U r s u s a r c t o s L . . Can. W i l d l . S e r . R e p t . S e r . No. 34, O t t a w a . 86 pp. T  P e a r s o n , A.M. and J.W. N o l a n . 1976. The e c o l o g y o f t h e g r i z z l y b e a r ( U r s u s a r c t o s 1.) i n J a s p e r N a t i o n a l P a r k — Report f o r 1975. U n p u E l . CWS R e p t . No. 25-76. 15 pp. P e t e r s , R.P. and L.D. Mech. 1975. S c e n t m a r k i n g A m e r i c a n S c i e n t i s t 63: 628-637.  i n wolves.  P i e k e l e k , W. and T.S. B u r t o n . . 1975. A b l a c k b e a r p o p u l a t i o n study i n northern C a l i f o r n i a . C a l i f . F i s h a n d Game 6 1 ( 1 ) : 4-2 5. P o e l k e r , R . J . and H.D- H a r t w e l l . 1973. B l a c k b e a r o f 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 . 14, O l y m p i a , Washington. 180 pp.  No.  Pyke, G. H. , H. R. . P u l l i a m , and E . L . C h a r n o v . 1977. O p t i m a l f o r a g i n g : a s e l e c t i v e review o f t h e o r y and t e s t s . Quart. Rev. B i o l . 52 ( 2 ) : 137-154. R e y n o l d s , H. and J . J . Beecham. 1978. Home r a n g e a c t i v i t i e s and r e p r o d u c t i o n o f b l a c k b e a r s i n w e s t - c e n t r a l Idaho. I n C . J . M a r t i n k a , e d . 4 t h I n t e r n a t i o n a l c o n f . on b e a r r e s e a r c h and management. K a l i s p e l l , Montana. R o c h e l l e , J . P e r s . Comm. W i l d l i f e Ltd., Centralia, Washington. Roth,  biologist,  Weyerhaeuser  H. 0. I n p r e s s . D e f e c a t i o n r a t e s o f c a p t i v e brown b e a r s . I n P r o c e e d i n g s o f t h e 4 t h I n t e r n a t i o n a l C o n f e r e n c e on Bear R e s e a r c h and Management. . C . J . M a r t i n k a , e d . K a l i s p e l l , Montana, 1977.  R o g e r s , L . L . 1976. E f f e c t s o f mast and b e r r y c r o p f a i l u r e s on s u r v i v a l , g r o w t h , and r e p r o d u c t i v e s u c c e s s o f b l a c k b e a r s . T r a n s . N. Am. W i l d l . and Nat. Res. Conf. 41: 431-438. R o g e r s , L . L . 1977. S o c i a l r e l a t i o n s h i p s , movements, and p o p u l a t i o n dynamics o f b l a c k bears i n n o r t h e a s t e r n Minnesota. U n p u b l . Ph.D. T h e s i s . U n i v e r s i t y o f Minnesota. 203 pp. R u d n a i , J.A. 1973. The s o c i a l l i f e o f t h e l i o n . M e d i c a l and T e c h n i c a l P u b l i s h i n g Company L t d . , London. 122 pp. R u s s e l l , D. 1974. G r i z z l y b e a r i n Knight I n l e t . B.C. F i s h Nanaimo. 92 pp.  - mountain goat i n v e s t i g a t i o n s and W i l d l i f e Branch P r o j e c t ,  127  R u s s e l l , R-H. 1971. Summer a n d autumn f o o d h a b i t s o f i s l a n d and m a i n l a n d p o p u l a t i o n s o f p o l a r b e a r s - a c o m p a r a t i v e study. M.Sc. T h e s i s , U n i v . o f A l b e r t a , Edmonton. v i + 87 pp. R u s s e l l , R.H., J.W. N o l a n , N.A. Woody, and 6. A n d e r s o n . Ecology o f g r i z z l y bears i n J a s p e r N a t i o n a l Park. prep.).  1978. (In  S c h a f f e r , S.C. 1971. Some 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 g r i z z l y b e a r s and b l a c k b e a r s o f t h e Apgar M o u n t a i n s i n G l a c i e r N a t i o n a l P a r k , Montana. U n p u b l . M.Sc. T h e s i s , U n i v . o f Montana. i x + 133 pp. S c h o e n e r , T. 1971. T h e o r y o f f e e d i n g E c o l . S y s t . 11: 369-404.  strategies.  Ann.  S h a c k l e t o n , D.M. p e r s . comm. A s s t . P r o f e s s o r , Dept. S c i e n c e , UBC, V a n c o u v e r , V6T 1W5.  Rev.  Animal  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 t h e b e h a v i o r a l sciences. M c G r a w - H i l l , New Y o r k . x v i i + 312 pp. S m i t h , B. 1978. P r e l i m i n a r y i n v e s t i g a t i o n s i n t o b l a c k and g r i z z l y bear responses t o c o a s t a l l o g g i n g . Unpubl. B.Sc. T h e s i s , Simon F r a s e r U n i v e r s i t y , V a n c o u v e r . 70 pp. S m i t h , G.W. p e r s . comm. with and c o n s e r v a t i o n o f f i c e r , Nanaimo.  photographs. Wildlife technician B.C. F i s h and W i l d l i f e B r a n c h ,  S t o n o r o v , D. and A.W. S t o k e s . 1972. S o c i a l b e h a v i o u r o f t h e A l a s k a brown b e a r . Pp. 232-242 i n S. H e r r e r o , e d . B e a r s t h e i r b i o l o g y and management. IUCN New S e r i e s No. 23, Morges, S w i t z e r l a n d . S t o r e r , T . I . and L . P . T e v i s . 1955. C a l i f o r n i a g r i z z l y . U n i v e r s i t y o f C a l i f o r n i a P r e s s , B e r k e l e y a n d Cambridge U n i v e r s i t y P r e s s , London. 335 pp. T i s c h , E . 1961. S e a s o n a l f o o d h a b i t s o f t h e b l a c k b e a r i n t h e W h i t e f i s h Range o f n o r t h e r n Montana. M.Sc. T h e s i s , U n i v . o f Montana, M i s s o u l a , i x + 108 pp.. Van  S o e s t , P . J . and R.H. Wine. 1967. Use o f d e t e r g e n t s i n t h e analysis of fibrous feeds. I V . The d e t e r m i n a t i o n o f p l a n t c e l l wall constituents. J . A s s n . O f f . A n a l . Chem. 50: 50.  Waldern, D.E. 1971. A r a p i d m i c r o d i g e s t i o n p r o c e d u r e f o r n e u t r a l and a c i d d e t e r g e n t f i b r e . Can. J . Anim. S c i . 51: 67. W e i s s , J . p e r s . comm. P l a n t e c o l o g i s t and a e r i a l p h o t o g r a m i t i s t . B.C. F o r e s t P r o d u c t s L t d . , 1050 W. P e n d e r S t . , V a n c o u v e r , B.C.  128  W h i t t e m o r e , C.T. nutrition.  and F.W.H. E l s l e y . 1976. Practical pig Farming P r e s s L t d . , S u f f o l k . 0 + 90 pp.  W r i g h t , W.H. 1909. New Y o r k . 268  The pp.  grizzly  bear.  Charles Scribner's  Z u n i n o , F. and S. H e r r e r o . 1972. The s t a t u s (Orsus a r c t o s ) i n Abruzzo N a t i o n a l Park, B i o l . Cons. .4: 263-272.  of t h e brown Italy, 1971.  Sons, bear  Appendix  I.  L a t i n names f o r common names u s e d i n ( i n the order i n which they appear)  Chapter  1  grizzly  bear  black  bear  Chapter salmon  Ursus  arctos  Ursus  americanus  3 -  sp.  Onaorhynchus  salmonberry  Rubus  sedge  Carex s p .  elk  Cervus  canadensis  beaver  Castor  canadensis  marten  Martes  americana  Chapter  ground  spectabilis  4  squirrels  sp.  Spermophilus sp.  mongooses  Herpestes  badger  Taxidea  taxus  lion  Panthera  leo  amabilis western sitka red  the  fir hemlock  spruce  pine  amabilis  Tsuga  heterophylla  Picea  sitchensis  Thuja  cedar  lodgepole  Abies  plicata  Pinus Canis  wol f  contorta lupus  ChaDter 5 i  polar  bear  Chapter  Ursus  maritimus  6  ladyfern  Athyrium  s p i n y wood  fern  Dryopteris  huckleberry  Vacciniwn  devi1 s  Oplopanax  1  club  filix-femina austriaca sp. horridum  skunk  cabbage  Lysiahitum  stink  currant  Ribes  tall  blue  alaskan  huckleberry  huckleberry  americanum  braoteosum  Vaocinium  ovali.foti.um  Vaocinium  alaskaense  chum s a l m o n  Onaorhynchus  keta  pink  Onaorhynchus  gorbuscha  salmon  trout  Salmo  wasps  Vespidae  bees  Apidae  deer  fern  bracken oak  Blechnum  fern  coho  spiaant  Pteridium  fern  sword  alarki  aquilinum  Gymnocarpium  fern  Polystiahwn  salmon  munitum  Onaorhynchus  horsetai 1  Equisetum  sockeye  Onaorhynchus  salmon  swi ne lowbush  dryopteris kisutah  sp. nerka  Suidae blueberry  Himalayan  black  bear  Vaocinium Selenarctos  scoparium thibetanus  Appendix  A.  Plant  Coniferous  Tsuga  Deciduous  trees  amabilis  Pioea  i n the Ahnuhati and 1977  species  heterophylla  Abies Thuja  L i s t of s p e c i e s found w a t e r s h e d d u r i n g 1976  II.  Alnus  rubra  Alnus  sitahensis  trees  sinuata  Populus  trichoaarpa  Salix sp.  pliaata  Abies  lasioaarpa  Picea  engelmannii  Tsuga  mertensiana  Chamaeayparis  Acer  maorophyllum  Pyrus  fusoa  Rubus  speotabilis  nootkatensis  Taxus b r e v i f o l i a  Upright  shrubs  Vacainium  alaskaense  Vaooinium  ovalifolium  Vacainium  parvifolium  Vaooinium Cornus  stolonifera  Oplopanax Menziesia Spiraea  membranaoeum  horridum ferruginea douglasii  Rubus p a r v i f l o r u s Ribes  braoteosum  Sambuous  raoemosa  Sorbus  sitahensis  Cladothamnus Viburnum Rosa s p .  pyrolifiorus  edule  132  Creeping  Rubus  shrubs  pedatus  Vacainium  uliginosum  Vacainium  ovatum  Gaultheria  Cassiope  mertensiana  Phyllodoce  glanduliflora  Phyllodoce  empetriformis  hispidula  Herbs  Urtica  dioica  Heracleum  Oxyria  digyna  Oenanthe  sarmentosa  Osmorhiza  chilensis  Stellaria Aotaea  longifolia  rubra  Aquilegia Coptis  Cornus  formosa sibirica  Trautvetteria Achlys  oarolinensis formosa  Saxifraga  ferruginea  Tellima  uniflora •  Pyrola  secunda sp.  Pyrola  triphylla  Dioentra  canadensis  Pyrola  asplenifolia  Montia  lanatum  grandiflorum  Chimaphila Monotropa  uni f l o r a  Trientalis aratica Menyanthes  trifoliata  Nephrophyllidium  Tiarella unifoliata  Romanzoffia  Tiarella'  Prunella  trifoliata  umbellata  crista-galli sitahensis  vulgaris  Heuchera  micrantha  Castillega  Aruncus  Sylvester  Mimulus  tilingii  maorophyllum  Mimulus  guttatus  Mimulus  lewisii  Geum  'Potentilla  glandulosa  Sanguisorba  sitahensis  miniata  Penstemon  davidsonii  Penstemon  serrulatus  Viola  palustre  Viola  glabella  Galium  boreale  alpina  Galium  cymosum  Ciroaea Epilobium Epilobium Epilobium  angustifolium alpinum watsonii  Valeriana  scouleri  Campanula r o t u n d i f o l i a Pinguicula  vulgaris  Achillea  millefolium  Smilacina  racemosa stellata  Anaphalis  margaritaoea  Smilacina  Prenanthes  alata  Streptopus  triangularis  Streptopus  Senecio Lysiahitum  americanum  Clintonia  uniflora  Erythronium Lilium  revolutum  columbianum  Maianthemum sp.  Angelica Aster  dilatum  Fragaria  sp.  Car ex  Trillium  ovatum  Veratrum  viride  Corallorhiza Goody era  maculata  oblongifolia  Habenaria Lupinus  sp.  roseus  saccata  Frittilaria sp.  sp.  Disporum  amplexifolius  sp.  Sedum s p .  sitchensis  Carex  miaroptera  sp.  Equisetum  sp.  Lyoopodium  Selaginella sp.  Ferns  Adiantum Athyrium  pedatum - fi  Blechnwn Cryptogramma  lix-femina  spicant arispa  Dryopteris Gymnocarpium  austriaoa dryopteris  Polystichum  muni turn  Pteridium  aquilinum  B.  Bird  harlequin common  duck  hawk  eagle  American blue  blue  heron  pigeon  unidentified  rufous  hummingbird  belted  kingfisher  common  flicker  yellow  bellied  sapsucker  green  herodias oanus  Columba fas  data  Cypseloides  niger  Selasphorus  rufus  Megaceryle  aloyon  creeper  dipper wren  robin thrush  sp.  (red race)Sphyrapicus  varius  Taahycineta  chickadee chickadee  nuthatch  villosus  sordidulus  Cyanocitta crow  ruber  Nuttallornis.borealis  swallow  backed  red-breasted  varied  Ardea  Corvus Corvus  black-capped  winter  obscurus  leuaurus  Contopus  raven  brown  Lagopus  pewee"  jay  Northwestern  chestnut  sparverius  Dendrooopos  flycatcher  wood  Steller's  leucocephalus  Colaptes  woodpecker  olive-sided  violet  jamaicensis  owl  swift  western  Buteo  Larus  band-tailed  hairy  merganser  Dendrogapus ptarmigan  gull  black  Mevgus  Falco  grouse  great  histrionicus  Haliaeetus  kestrel  white-tailed  mew  Histrionicus  merganser  red-tailed bald  species  thalassina stelleri  oaurinus aorax  Varus a t r i a a p i l l u s Parus  rufesaens  Sitta  canadensis  Certhia  familiaris  Cinelus  mexicanus  Troglodytes  troglodytes  Turdus  migratorius  Ixoreus  naevius  Swainson's hermit  thrush  thrush  ruby-crowned  kinglet  golden-crowned cedar  kinglet  Regulus  satrapa cedrorum  pusilla  warbler  tanager  Dendroica Piranga  grosbeak  nigrescens. ludoviciana  Pheucticus  melanocephalus  pine  siskin  Spinus pinus  song  sparrow  Melospiza  Oregon  junco  C.  Douglas  species  shrew  Sovex vagrans  squirrel  Tamiasoiurus  beaver  Castor  white-footed  deer  mouse  wolf  douglasi  canadensis  Peromyscus  maniculatus  Canis lupus  American grizzly  melodia  Junco oreganus  Animal  wandering  black bear  bear  Ursus americanus Ursus  arctos  marten  Martes americana  wolverine  Gulo  cougar  Felis  blacktail mountain  :  Dendroica auduboni  black-throated gray  black-headed  guttata  Wilsonia  warbler  western  Hyloeichla  Bombycilia  warbler  Audubon  ustulata  Regulus calendula^  waxwing  Wilson's  Hylocichla  deer goat  luscus concolor  Odocoileus  hemionus  Orearrmus americanus  Appendix I I I .  Seasonal  s h i f t s i n use of foods by Ahnuhati  black and g r i z z l y bears durir.g 1976.  H A Y  0 16-•31 •31  1-15 Food Species  U  V  IV  %  0.08  1.9  1.5  tr  ladyfern and spiny wood fern  0.93  27.6  25.8  34.8  0.77  6.2  4.8  sedge  0.60  18.0  10.8  14.6  0.57  9.2  4.3  skunk cabbage  0.13  4.0  0.52  0.7  leaves o f huckleberry  0.8  12.3  9.9  13.0  0.62  7.3  4.5  shoots o f salmonberry  0.73  11.0  14.8  1.0  50.8  leaves o f d e v i l ' s c l u b  0.87  13.9  18.7  0.85 0.11  V  IV  %  horsetail  0.15 16.0  cow parsnip chocolate  E -30 16--30  f  f  N  1-15 f  V  IV  %  IV  %  21.0  16.0  22.0  20.0  13.4  18.0  0.22  11.0  2-3  3.0  5.4  0.22  2.2  0.49  50.8  61.4  0.89  25.0  22.3  31.0  21.2  18.0  21.7  0.89  20.0  17.8  25.0  1.2  1.3  tr  tr  tr  0.11  0.77  0.09  tr  f  V  6.0  0.78  5.2  1.7  1.0  lily  b e r r i e s of• huckleberry b e r r i e s o f salmonberry berries of d e v i l ' s club berries of stink currant other b e r r i e s ants wasps and bees other i n s e c t s salmon manma1 h a i r  tr  tr.  sand, g r a v e l , o r d i r t unknown Sample s i z e (H) =  0.33 15  6.6  2.2  2.9  tr  13 OJ  cn  Appendix I I ! .  Seasonal s h i f t s i n use o f foods by Ahnuhati  J U L  A  Y 16--31 -31  1 -15 Food s p e c i e s horsetail l a d y f e r n and spiny wood f e r n  f  V  . . . .  . . . .  0.8  28.0  black and g r i z z l y bears during 1976 (cont'd).  IV  22.4  %  26.0  f  0.89  V  25.0  IV  22.3  32.0  0.8  14.0  11.2  13.0  0.66  17.5  11.6  17.0  skunk cabbage  0.4  8.0  3.2  4.0  0.56  4.0  2.2  3.0  tr  shoots o f salmonberry  1.0  1 eaves o f d e v i l ' s c l u b  1.0  cow  7.0  1.5  2.0  tr  1.5  0.5  1.0  37.0  28.9  41.0  1.5  17.0  17.0  20.0  0.22  32.0  32.0  37.0  0.33  0.78  2.8 1.11 tr  lily  b e r r i e s o f huckleberry b e r r i e s o f salmonberry  U  S  T 16-31  IV  i  6.5  11.0  0.2  tr  0.24  tr  tr  tr  f  V  0.5  1.3  tr  tr  0.25  IV .  tr 1.3  0.63  % 1.0  tr  tr  tr  tr  0.33  tr  •  —— — —  -----  parsnip  chocolate  G  tr 0.22  0.22  11.7  0.8  tr  tr  V  .  0.56  0.33  tr  tr  f  %  sedge leaves o f huckleberry  U  11-•15  . . . .  0.56  2.5  4.5  0.78 0.66  4.0  berries of d e v i l ' s club  tr  0.5  tr  tr  berries of stink currant  tr  0.5  tr  tr  28.3 15.6 33.1  0.33  22.0 10.3 1.0  39.0 18.0 2.0  0.25  16.3  4.1  7.0  0.75  6.3  4.7  6.0  0.5  5.0  2.5  3.0  .—-  other berries 1.11  0.33  ants  0.37  tr  •  wasps and bees other i n s e c t s 36.0  0.44  salmon  16.0  28.0  1.0  63.0  63.0  80.0  0.2  6.0  3.0  4.0  mammal h a i r .  sand, g r a v e l , o r d i r t unknown Sample s i z e (N) =  0.2  0.22  21.0 10  tr  ,  Appendix I I I . (cont'd)  Seasonal  s h i f t s i n use of food by Ahnuhati  black and g r i z z l y bears during 1977.  H A Y 1-15 Food s p e c i e s horsetail  J U N E 16-31  IV  %  —  1-15 IV  1.0  % 1.0  lady fern and spiny wood f e r n  0.8  14.80  11.80  14.0  0.75  17.75  13.30 15.0  sedge  0.8  36.28  28.80  33.0  0.75  20.30  15.20 17.0  skunk cabbage  0.4  0.80  0.32  0.3  0.50  0.50  0.25 . 0.3  16-30 IV.  (  f  IV  0.14  0.86  0.12  0.2  0.2  10.8  2.2  0.71  13.10  9.30  12.0  0.2  3.2  6.4  1.0  0.86  47.60  40.90  53.0  1.0  53.6  53.1  76.0  0.29  1.29  0.37  0.5 0.4  26.4  10.6  15.0  0.6  5.0  3.0  4.0  0.2  0.2  0.04  tr  0.2  0.8  0.16  tr  3.0  leaves of h u c k l e b e r r y shoots of salmonberry  0.8  17.80  14.30  17.0  1.00  49.80  49.80 55.0  0.71  17.60  12.50  16.3  leaves of d e v i l ' s c l u b  1.0  30.80  30.80  36.0  1.00  11.75  11.75 13.0  0.71  18.90  13.40  18.3  0.14. 0.29  0.04  tr  b e r r i e s of salmonberry  0.14  0.14  0.02  tr  b e r r i e s of d e v i l ' s c l u b  0.14  0.14  0.02  tr  0.14  0.14  0.02  tr  cow parsnip chocolate  lily  b e r r i e s o f huckleberry  b e r r i e s of s t i n k c u r r a n t other b e r r i e s ants wasps and bees other i n s e c t s salmon mammal h a i r sand, g r a v e l , o r d i r t unknown Sample s i z e (N) =  Appendix I I I . (Cont'd)  Seasonal s h i f t s i n use o f food by Ahnuhati  black and g r i z z l y bears during 1977.  A  J U L Y IV  Food s p e c i e s  0. 05  0. 44  tr  16--31  V  IV  * tr  IV  V  f  2  %  0.17  0.33  0.06  tr  0.2  0.4  0.08  0.2  0.4  0.08  tr  5.0  2.0  1.0  2.0  0.4  19.6  7.80  12.0  0.5  11.0  5.8  12.0  8.6  5.20  8.0  3.8  3.80  6.0  sedge  0..44  9. 33  4. 10  6.0  0.67  14.30  9.50  11.0  skunk cabbage  0..22  5..11  1. 12  2.0  0.50  2.50  0.13  tr  leaves of h u c k l e b e r r y  0.,33  2..80  0. 92  1.0  0.83  4.17  3.50  4.0  shouts o f salmonberry  0..78  7..30  57. 00  9.0  0.33  0.83  0.27  tr  leaves o f d e v i l ' s c l u b  0..11  2..80  3. 10  tr  0.17  0.33  0.06  tr  0,.11  1..90  2.,10  tr  berries of huckleberry  0.89  36.00  32.00  50.0  b e r r i e s o f salmonberry  0.89  18.00  16.40  26.0  berries o f d e v i l ' s club  0.44  1.89  0.83  1.0  berries of stink currant  0.11  0.33  0.04  tr  other b e r r i e s  0.11  0.30  0.33  tr  0.44  1.11  0.49  chocolate  T  IV  0. 33  cow parsnip  S  V  l a d y f e r n and spiny wood f e r n  tr  U  f  0. 11  0. 15  -  f  horsetail  0. 44  G  1-15  16--31  1-15  U  0.33  lily 1.00 1.00 0.83  0.50  38.20 22.80 14.17  0.2  0.17  38.70 22.80 11.80  • 0.6 1.0  1.0  44.0  0.2  26.0  1.0  14.0  0.8  14.2 1.8 7.2 33.4  14.20 0.36 7.20 26.70  —  0.5  0.5  0.25  tr  0.5  0.5  0.25  tr  1.0  0.5  0.5  2.50  5.0  11.0  0.5  3.5  1.80  4.0  0.5  3.5  1.80  4.0  0.5  2.5  1.30  3.0  0.5  50.0  25.00  0.5  21.0  4.0  _  tr  ants wasps and bees  tr  other i n s e c t s salmon mammal h a i r  0.11  7.56  0.83  0.60  sand, g r a v e l , o r d i r t  0.11  0.89  0.10  tr  unknown  0.22  0.22  0.05  tr  Sample s i z e (N) =  9  0.33  0.33  --•--  ——  0.21  tr  0.20  9.4  tr  tr  tr  0.11  -—  .. 1.90  3.0  —  '  -—- — — 6 —  0.5 —  0.21 10.50  0.21  2 CD  Appendix I I I .  Seasonal s h i f t s i n use o f food by Ahnuhati  (Cont'd)  S  E  1-15 Food s p e c i e s  f  IV  V  P T E  %  M  B  E  black and g r i z z l y bears during 1977.  0  R 16-30  f  V  IV  %  c T  0  B  E  R 16- 31  1-15 f  V  IV  f  %  —'.  IV  V  %  0.8  tr  tr  tr  0.8  2.0  1.60  2.0  0.5  1.0  0.5  tr  l a d y f e r n and spiny wood f e r n  0.6  1.8  1.08  1.0'  0.8  8.6  6.90  7.0  1.0  35.5  35.5  52.0  sedge  0.2  5.2  1.04  1.0  0.8  6.4  5.12  5.0  0.5  3.0  1.5  2.0  tr  0.4  0.2  0.08  tr  1.0  1.0  4.6  4.60  5.0  tr  0.4  0.8  0.32  tr  horsetail  s k u n k cabbage  0.4  tr  tr  leaves o f huckleberry  0.8  1.2  0.95  shoots o f salmonberry  0.4  tr  tr  ....  leaves of d e v i l ' s c l u b  ----  cow parsnip chocolate  lily  , —-  berries of huckleberry  0.2  0.8  0.16  tr  b e r r i e s o f salmonberry  0.6  0.4  0.24  tr  .... 0.4  0.2  0.08  tr  berries of d e v i l ' s club  0.2  tr  tr  tr  b e r r i e s of s t i n k c u r r a n t  0.2  0.67  0.13  tr  -  -  0.2  0.78  0.28  tr  .—— C.4  wasps and bees  1.0  61.20  61.20  0.66  1.0  0.2  0.08  tr  62.4  62.40  66.0  .... '  other i n s e c t s salmon mammal h a i r sand, g r a v e l , o r d i r t unknown Sample s i z e (N) =  1.0 0.2  ....  0.2  28.60  28.60  0.31  1.0  0.20  1.0 12.5  12.50  tr  •  13.0  tr  tr 5  • .... -—  .  other berries ants  —..  5  0  —. .0.5  33.5  16.8  25.0  -  •  0.5  12.5  6.3  9.0  •  0.5  12.5  6.3  9.0  -—  •  .0.5  14.5  7.3  11.0  2  o  141  Appendix  IV.  Abbreviations  Nutrient  values of  -  DO  =  d r y , open  canopy  DC  =  dry, c l o s e d  WO  =  wet, open  WC  =  wet, c l o s e d  pdm  =  percent dry  CHO  =  carbohydrates  canopy  canopy canopy matter  bear  foods  A)  Data  H  D e v i l ' s c l u b : (1) leaves  (eaten).  Sample  Site  r e f . no.  characteristics  47 54 31  DO DC WD  X J  76 74 91 73  WC DO WD DC  .  X A  99 16 18  00 WC WD  X A) M  Devil's club : 48 33 29  83 67 95 62 X  (2)  (3)  (4)  (5)  Crude p r o t e i n  Soluble  (6)  Crude f a t  (7)  Crude f i b e r  (8)  Gross energy  Minerals  Fibre:protein  Moisture content  (pdm)  CHO  (pdm)  33.38 25.44 24.75  17.82 22.14 33.71  27.85  (pdm)  .  (pdm)  (Kcals/g)  (pdm)  ratio  0.90 1.44 1.16  88.98 93.21 90.79  (*)  4.19 5.98' 2.19  30.06 36.64 28.62  4.505  9.21 • 9.80 10.73  24:55  4.12  31.77  4.505  9.90  1.17  91.00  14.25 16.25 9.63 18.88  38.81 39.28 53.98 33.62  2.31 2.06 2.07 1.89  36.11 32.19 26.24 36.96(34.0)  4.506 4.383  8.52 10.22 8.08 8.65  53 98 73 96  85.50 87.56 80.27 88.29  14.80  41.43  2.08  32.83  4.440  8.87  2.30  85.42  11.00 14.13 9.94  24.28 40.87 • 47.68  2.84 2.09 2.27  51.98 32.29 29.76  4.300 4.405  9.90 10.62 10.35  4.73 2.29 2.99  82.37 84.13 78.60  11.69  37.61  2.40  38.01  4.3525  10.29  3.34  81.60  6.19 3.32 3.38  27.19 28.38 30.06  3.79 5.37 3.34  58.26 59.64 59.91  4.661  4.57 3.29 3.31  9.40 17.96 17.72  76.80 .75.08 92.05  4.30  28.54  4.17  59.27  4.661  3.72  15.03  81.30  3.69 4.25 3.44 3.69  35.65 29.03 30.10 43.23  3.17 3.56 3.97 2.42  54.64 60.43 60.17 47.70  3.77  34.50  3.28  55.70  -  (2) s t a l k ( n o t e a t e n ) . DO DC WD  X J  0)  DO DC WC WD  ' 4.60  4.60  2.85 2.73 2.32 2.96  14.80 14.20 17.49 12.93  63.10  2.72  14.86  63.10  B)  Vaccinium:  M  49 50 51 52  (4)  berries  DO WC WC DC  105 103 no  WO DO al. ov.  X A  113 114 115 115  WDDO DC DO  X C) M  Combined  sample  27 46 30  of  Ladyfern DO WC WD  X J  63 78 77 94  DO DC WC WD  X JL  24  (5)  (6)  (7)  (8)  (2)  18.94 19.13 15.69 20.32  43.71 36.69 44.16 36.31  2.13 3.30 2.80 2.45  31.79 36.74 34.08 37.37  4.89 4.89  3.43 4.14 3.27 3.55  1.68 1.92 2.17 1.84  82.00 80.00 84.00 82.00  18.50  40.22  2.67  35.00  4.89  3.60  1.90  82.00  8.56 8.63 11.00 6.13  61.50 63.02 61.55 67.25  12.79 10.71 10.13 10.32  14.80 15.32 14.99(15.26) 14.00  2.35 2.32 2.33 2.30  1.73 1.78 1.36 2.28  88.10 85.50 87.30 89.50  8.58  63.30  10.99  14.78  5.00  2.33  1.79  87.60  5.94 6.50 6.44 7.75  65.67 65.16 64.84 64.83  11.19(8.93) 10.11 11.57 10.33  14.88(15.43) 15.83 14.70 14.98  4.90  2.32(2.36) 2.40 2.45 2.11  2.50 2.44 2.28 1.90  90.70 87.70 88.20 88.10  6.66  65.13  10.80  15.10  4.90  2.32  2.28  88.70  37.56 36.30 32.44  29.47 9.29 19.03  1.70 2.15 1.86  22.10(22.71 ) 44.40 36.38  4.52  10.90 7.86 10.29  0.59 1.22 1.12  91.40 92.50 92.05  35.43  19.26  1.90  34.29  4.52  9.68  0.977  92.00  17.00 18.06 22.25 22.31  36.58 37.24 35.74 33.77  1.49 2.27(2.82) 1.79 4.65  35.85 33,58 33.10 32.50  4.60  9.08 8.85(8.94) 7.12 6.77  2.11 1.86 1.49 1.46  84.58 82.17 83.20 80.98  19.90  35.80  2.55  33.76  4.60  7.96  1.73  82.70  22.56  24.59  2.29  42.44  4.66  8.02  1.90  (.eaten).  I U  (3)  (4)  (1)  DO  and  Spiny  wood  5.00  4.90  fern.  (2)  (3)  (4)  (5)  17.06 14.00 16.19 16.13  23.17 39.71 35.18 25.20  2.12 2.79 1.74 2.46(2.60)  43.03 34.29 40.02 45.29  4.50  15.85  30.81  2.28  40.66  4.50  25.69 26.19 29.69 16.94  39.86 38.12 39.41 50.17  4.40 1.32 2.25 1.24  23.56 29.09(31.17) 23.06 25.72  24.63  41.89  2.30  •25.35  14.75 14.44 15.13 11.94  30.25 35.04 32.45 27.11  1.58 1.49 1.06 1.70  48.78 43.20 46.02 54.46  14.07  31.20  1.46  48.10  8.56 9.13 13.38 7.00  32.23 31.11 26.27 32.72  2.39(2.30) 1.53 ' 1.48 1.14(1.51)  9.52  30.60  21.10  0) A  97 103 102 5  DO DC WC WO  X D)  Salmonberry: (1) shoots ( e a t e n ) .  M  55. 57 53 44  DO DC . WC WD  X J  65 87 81 89  DO DC WC WD  X A  4 12 5 8  DO DC WC . WD  X S °) M  38  T  ' (6)  (8)  (7)  11.89(11.71)2.52 9.21(9.22) 2.45 6.87 2.47 10.95 2.80  76.80 79.75 71.20 75.70  9.73  2.56  75.90  6.49 5.28 5.59(5.77) 5.93  0.92 1.11 0.78 1.52  91.17 92.33 93.21 88.36  5.83  1.08  91.27  4.64 5.83 5.34 4.79  3.30 2.99 3.04 4.56  65.35 81.16  4.39  5.15  3.50  73.30  51.93 53.41 53.09(51.72) 55.15  7.03 4.44 7.17 4.40  4.89 4.82 5.78(5.79) 3.99  6.10 5.85 3.97 7.88  66.74 72.59 63.03  1.64  53.40  5.76  4.87  5.95  67.50  31.88  2.26(2.36)  34.94  4.35  9.82  1.66  4.88 5.06 4.63  25.61 24.47 23.79  1.24 1.35 2.32(1.91)  66,87 67.44 68.12  4.80  1.40 1.68 1.14  13.70 13.32 14.70  4.86  24.60  1.64  67.50  4.80  1.41  13.90  4.59  -----  4.59 4.39  '  Salmonberry: (2) s t a l k (not e a t e n ) . 43 56 61 X  DO DC WC  .  59.16 61.42 62.41 61.00  84 72 66  DO DC WC  X 100 11 19 40  DO DC WC WD  X  (1)  (2)  4.00 3.94 4.75  27.79 27.21 27.28  0.89 0.39 1.30  65 .36 66 .72 65 .68  3.17  27.43  0.86  65.92  35.37 32.06 17.97 34.20  1.43 1.28 0.78(1.45) 1.69  58 .15 61 .31 64 .79 58.74(59.44)  33.88  1.47  59,.40  19.56 22.69 25.80 . 18.00  47.61 38.54 33.65 50.66  1.82 1.96(2.04) 2.61 0.69(1.79)  25.00 29..86 30..94 26..97  21.50  42.62  1.77  28..19  13.69 19.81 22.63 15.38  51.90 41.63 36.33 49.53  2.59 2.33 2.57 2.73  20. 35 28. 66 31. 39 26. 71  19.13  44.85  2.56  26. 78  11.13 11.80(14.0) 22.38 12.50  36.57 43.71 38.89 53.88  3.13 3.21(3.61) 2.80 2.36  14.45  43.26  2.88  3.56 3.69(3.94) 14.06 3.56 3.60  (3)  (4)  (5)  .  (7)  4.60  1.96(2.13) 1.74 ' 0.99  16.34 16.90 13.80  59.79 61.69 61.20  4.60  1.56  15.68  60.90  1.49 1.66 2.40 1.81  16.33 16.60 4.60 16.50  43.28 46.65 60.86 51.60  1.65  16.50  47.20  4.47  6.01 6.95 7.00 3.68  1.27 1.32 1.20 1.50  83.45 88.01 81.26 85.77  4.49  5.91  1.33  84.60  6.47 7.57 7.08 5.65  1.10 1.45 1.39 1.74  71.80 .78.15 78.92 67.00  4.47  6.69  1.42  74.00  43. 75 32. 79 31. 04 24. 62  4.40 4.30 4.46  5.42 8.49 9.00 6.64  3.93 2.78 1.39 1.97  64.42 70.80 77.37 65.77  33. 10  4.39  7.39  2.52  69.60  4.47  4.47  (6)  •  (8)  Salmonberry: (3) leaves ( p o s s i b l y e a t e n ) . 45 26 59 28  DO DC WC WD  X 93 75 79 92  DO DC WC WD  X 14 1 3 41 X  DO DC WC WD  4.50  4.47  0) D) J  107 109  WO DO  X A  (2)  (3)  (4)  10..75 12 .94  58.81 57.29  8.95 8.28(7.32)  18.60 18.43(18.60)  11..85  58.10  8.62  18.52  12..88  45.66  13.82  25.52  (5)  (6)  (7)  (8)  4.94  2.89 3.06  1.73 1.42  85.20 86.50  4.94  2.93  1.58  85.90  2.12  1.98  83.50  Salmonberry: (4) b e r r i e s .  112  WO  E)  Skunk cabbage : (1) r o o t (eaten).  M  34  WC  15..25  35.52  2.42  31.14  4.098  15.67(16.10)  2 .04  95..80  J  85 '70  WD' WC  13..13 18..33  36.15 29.72  2.68 1.53(1,.97)  33.62(33.10) 32.52  3.90  14.42 17.90(18.30)  2 .56 1 .77  93..16 95,,20  15..73  32.94  2.105  33.07  3.90  16.16  2,.17  94,,20  X JL  23  WD  13..88  45.68  2.070  25.71  4.10  12.66  1,.85  A  105 7  WO WC  20..56 20: ,38 .  28.51 19.17  1.280 1.56(1,.58)  32.42 35.75  6.57  17.23 23.14  1,.58 1,.75  88,,21 91.,23  20. 47  23.84  1.42  34.10  6.57  20.19  1..67  89.,70  20.69 24. 63 19.38  43.99 40.74 46.30  2.18 3.25 3.0(3.79)  19.45 18.81 19.59(20.32)  4.143  13.69 12.57 11.73  0..94 0,.764 .1,.01  21. 57  43.68  2.81  19.28  4.143  12.66  0.905  4.46  11.89  1..04  92.,50  X S  22 37 21  WC WC WC  X E)  Skunk cabbage : (2) leaves (not e a t e n ) .  M  35  WC  23. 56  37.08  2.90  24.57  J  82 71  WC WD  20. 69 12. 50  34.08 43.60  2.50 1.83  22.55 26.12  3.90  20.18 15.95  1.,09 2..01  93. 84 93. 50  16. 60  ' 38.84  2.17  24.34  3.90  18.07  1.,55 .  93. 70  X  .  15 13  A  WC WD  X  (1)  (2)  (4)  (5)  19.10 18.00  30.80 27.70  3.02 3.51  28.79 32.48  -.00 1.89  18.26 18.32  1.51 1.80  90.84 91.55  18.55  29.30  3.27  30.64  3.95  18.30  1.66  91.20  2.63  1.16  69.32  8.62  0.91  71.80  5.63  1.04  70.60  5.67  2.24  72.61  5.40  2.61  8.47  2.26  Mg M J  A  .  M  Ca  K  0.13 0.143  0.68 0.60  7.2 5.2  0.11 0.14  0.93 1.0  8.6 10.4  leaves  0.174 0.167  1.1 1.24  6.8 7.1  WO WD  21.06 26.81  44.02 39.52  2.07 1.61  30.22 24.44  4.680 4.55  23.94  41.80  1.84  27.33  4.62  WC WC  root  WC WC  root  WC WC  root  1 eaves leaves  (7)  (8)  Sedge 32 . 25 X  J  88  WD  16.81  35.23  1.16(2.01)  37.69  4.62  JL  39  WO  14.25  41.65  1.57  37.13  4.67  A  42  WO  15.30  40.46  1.12(2.20)  34.65  4.59  G)  Two female coho  A  (6)  Skunk cabbage: (3) m i n e r a l s .  E)  F)  (3)  A  72.04 74.31  A  73.20  7.95 7.95  4.95 6.19 5.60  •  71.16  4.886 5.109  10.84 10.84  62.57 78.02  4.998  10.84  70.30  Appendix  V.  Example o f d e t e r m i n a t i o n  o f net energy  from the energy  crude  protein  (pdm) Proximate 1.  2.  3.  a n a l y s i s o f two f e m a l e  cohos  showed:  reduce according d i g e s t i b i l i t y of gross energy (87.6 p e r c e n t ; Hamilton 1978). energy 1961).  lost  to urine  (0.9 kcal/g;  energy kcal/g  l o s t t o methane p r o d u c t i o n CHO; B r e i r e m 1 9 3 9 ) .  available  soluble carbohydrates (pdm)  i n coho  salmon.  crude fat (pdm)  73.2  7.95  5.6  (.876) 73.2 = 64.1  7.01  4.0  moi s t u r e content (pdm) 70.3  Kleiber 57.7  4.  caloric equivalents  5.  amount o f e n e r g y  6.  delete  7.  e f f i c i e n c i e s of conversion ( F i n g e r l i n g 1918).  values  8.  net energy  9.  caloric  o f 2.  and 3 .  p e r 100 g d r y  dry  1961).  present  density  pdm = p e r c e n t  (Kleiber  matter  (1.02 7.14 5.7  kcal/g  4.0  9.5  365.4  kcal  28.0  38.0  307.7  kcal  20.9  38.0  to f a t  matter  0.74 227.7 2.8  0.83 kcal  14.4  k c a l / g dry matter  0.93 35.0  o r 848 k c a l / k g wet  =  280.1  weight  kcal  Appendix V I .  Bear • ID No.  Species  19/23  B  Sex  M  Age  • 7.5  Body measures and i m m o b i l i z a t i o n p a r t i c u l a r s f o r bears captured i n the Ahnuhati  125  E  250  E  59  E  59  E  H99  G  11  M  165  1.3  M99  5.0  M99  4.8  M99  2.0  Sern.  180  no"  H99  4.0  113  M  sern.  205  14  E  M99  1.0  64  H  Sern.  130  M99 Sern.  second capture  20B  B  F  2  E  18  B  F  2  E  7P  G  F  21  G  F  21-A  G  H  20G  G  F  7C  G  F  4  B  M  E  = estimated  — — 0.5  — •— —  Relative dose (mg/kg) 0.034  •Sern.  —  Absolute dose (mg) 4.2  second capture 12/22  Drug used  Weight (kg)  144» 48  M  H  =  Induction (min) 8  Time ur.conc. (hrs) 3  Capt. date . - Scars  19/6/76  12  4.3  0.02  6  4.0  0.02  11  2.0  24/6/76  during 1976 and 1977.  Total length (cm)  no  2 fresh  Neck ' girth (cm)  Chest girth (cm)  Width 1 ear: (cm  68  176  168  —  32  2 fresh  6  —  no  142  53  no  12  35  4.5  no  140  55  115  12  0.04  5  1.1  no  179  160  103  19  1.8  34  4.5  no  151  59  107  19  0.07  4  0.75  no  83  30  46  10  2.0  9  —  no  134  35  83  14  4.0  0.03  4  2.5  no  197  61  114  22  60  1.3  10  4.0  no  130  38  79  10  measured  0.034 R  R  R  3.1  R  =  +  r e p e t i t i v e dose  Appendix  VII.  Frequency and p e r c e n t cover o f u n d e r s t o r y s p e c i e s i n 15 p l a n t communities  i n the Ahnuhati watershed,  1977.  P l a n t community type number 1_ Area Sample s i z e  2.14  2  3_  1.29  (N)  Plant species  Cavex sp. Species o f Graminae Lyaichitum americanum Viola palustre Oenanthe sarmentosa Splvighnum sp. Vaocinium alaskaense Vaocinium ovalifolium Vaocinium ovation Dryopteris oustriaca Blechnwn spicant Athyrium filix-femina Streptopus roseus Rubus pedatwn Corr.us canadensis Clintonia uniflora Menziesia ferruginea Tiarella uniflora Oplopanax korridum Rubus speotabilis Viola glabella Circaea alp-ina Ribes braoteosum Sambucus racemosa Trautvetteria -carolinsnsiB Streptopus amplexifolius  M5  11.2  26.8  76.5  2.02  32.5  0.81  30  H5  33  M7  3.9  M5-7  36  H6-7  12.6  1 f  PC  f  100 100 100 75 50 75 25 25  87 2.6 6 3.3 tr 49 tr tr  67 100 100 100 67 33 67 33  50 0.5  PC  6 16 69.3 2.7 0.7 9.3 tr tr  f  PC  f  PC  tr tr  55 13 40  tr 0  0  78 100 67 100 67 78  20.2 41.1 11.9 0.1 7.7 3.3  56 100 100 78 78 '67 56 44 11  1.2 12.2 2.4 0.1 2.0 1.8 2.0 0.6 0.1  f  PC  67 1.7 33 33 33 33 33 33  2.7 tr 23.3 tr 2.7 tr  33 100 100 100 100 67 33 67 33  tr 19 84.3 3 0.1 2.7 10 tr tr  67 43  50 0.8  67 0.7 33 t r  2 tr  50 t r 50 t r  33 t r 33 t r  2 tr  25 t r 25 t r  100 2.7 67 0.3 33 t r  frequency  PC  f  PC  f  PC  f  PC  f  14 t r 29 0.6  100 100 50 50 50  PC  f  PC  37.5 tr tr 2.0 2.0  33 33 33 67 33 33 67  tr 0.7 tr 10.3 1.3 tr 16  f  PC  f  PC  f  50.3  Est.  Est.  PC  f  PC  •  6.7 13.7 4.6 tr 5.0 1.1 9.5 0.6 5.0 0.3 0.1 0.1 8.3 29.6 26.4 0.2 1.5 4.2 0.3 0.8 0.1  50  0  75 t r  f  M5-6-7  3.8  11 t r 11 0.3  33 t r  PC  22 0.1 44 0.1 78 11 44 44  45 0.1 4.2 0.6  10  71 5.5 50 . 0  100 10.2 14 1.1  percent c o v e r  4.2 2.6 75.7 0.4 1.7 2.6 0.6 2.4 0.2  SN  0 2 5 0 tr 0  tr  14 29 71 100 14 71 57 86 29  tr 1.9 0.6 66.9 tr 0.6 3.7 2.3 2.3  100 3 100 t r 100 8  50 t r 100 t r 50 t r  trace  tr 0.4 tr 12.7 0.7 SN 8  0 0  50 t r 50 0.5  11 0.1 100 89 100 78 89 78 67 78 44  50 4.0  15  5 tr 0 15 • tr  100 100 100 67 100  10.3 4.3 34.3 1 tr  67 0.3 67 13 67 t r  8 25 55  15 18 2 38  8 0 8 .  SN = senescent  9.2 34 44.5 0.5 4.0 tr 4.2 6.5 tr  Appendix V I I .  (Cont'd).  P l a n t s p e c i e s cover In s u b a l p i n e meadow o f S u n k i s t V a l l e y , Ahnuhati  1977  (see F i g u r e 13).  Plant species  Carex  sp.  Valeriana  Frequency  •100 saouleri  Veratrwn  vivide  Sanguisorba  eitaheneis  Percent r o  v  e  r  37.8 8.4  100  25.5  25  7.0  Species of Gra'minae  50 .  2.5  Epilobium  alpinum  25  1.25  Bevacleum  lanatwn  75  3.5  25  1.0  25  6.0  75  5.0  75  0.63  25  1.25  25  0.63  Ribea  bracteoswn  Rubus  speatabilie  Athyviwi  filix-femina  Epilobium  anguQtifolium  Streptopue  roseus  TrautvetteHa Others  oarolinsneia  75  Figure  i  12:  Vegetation  c o m i t i e s  in  the  t  valleybottom  of  the  Ahnuhati  

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