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Qualities of individuals in a replacement stock of blue grouse Low, David J. 1975

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QUALITIES OF INDIVIDUALS I N A REPLACEMENT STOCK OF BLUE GROUSE by DAVID JAMES LOW B.Sc,  U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1971  A thesis  submitted i n p a r t i a l  the requirements  fulfillment of  f o r t h e degree o f  MASTER OF SCIENCE i n t h e Department o f ZOOLOGY  We a c c e p t t h i s required  t h e s i s as c o n f o r m i n g t o t h e  standard  THE UNIVERSITY OF B R I T I S H COLUMBIA A p r i l , 1975  In  presenting  this  thesis  an advanced degree at the L i b r a r y I  further  for  it  freely  this  thesis for  It  of  financial  <3O JLJOG. of  the  requirements  B r i t i s h Columbia, for  reference copying o f  X  I  I agree  not  for  that  and study. this  thesis  by the Head of my Department  gain s h a l l  B r i t i s h Columbia  2075 Wesbrook P l a c e Vancouver, Canada V6T 1W5  of  i s understood that copying or  written permission.  The U n i v e r s i t y  of  that permission for extensive  representatives.  Department  fulfilment  available  s c h o l a r l y purposes may be granted  by h i s of  the U n i v e r s i t y  s h a l l make  agree  in p a r t i a l  or  publication  be allowed without my  ABSTRACT  An e n t i r e b r e e d i n g p o p u l a t i o n o f b l u e grouse obscurus)  was  removed i n 1970  from an a r e a o f 900  (Dendraqapus acres of  second-growth f o r e s t l a n d on c e n t r a l Vancouver I s l a n d . p o p u l a t i o n had been d e c l i n i n g s i n c e 1954. i t s grouse p o p u l a t i o n i n t a c t was  The  A nearby area w i t h  r e t a i n e d as a c o n t r o l .  The  a r t i f i c i a l l y removed p o p u l a t i o n was  completely r e p l a c e d a t  the f i r s t b r e e d i n g season,  completely by  b i r d s , which i s evidence duals i n the s p r i n g .  - almost  yearling  f o r there being a surplus of  T h i s new  p o p u l a t i o n resembled  o r i g i n a l r e s i d e n t s and o t h e r grouse  i n surrounding  indivi-  the areas.  The use o f behaviour e v a l u a t i o n t e c h n i q u e s r e v e a l e d t h a t the r e c o l o n i z i n g p o p u l a t i o n was r e s i d e n t s had been.  The  evidence t h a t b l u e grouse territorial  behaviour.  as a g g r e s s i v e as the  removal experiment space themselves  original  gave s t r o n g out  through  ii  TABLE OF CONTENTS Page ABSTRACT  i  TABLE OF CONTENTS  i  L I S T OF TABLES  i i v  INTRODUCTION  1  MATERIALS AND METHODS  3  Population  analysis  3  Behaviour  4  a)  Mirror t e s t behaviour  4  b)  Grouse b e h a v i o u r r e c o r d e d by o b s e r v e r s in  the evenings  5  RESULTS  6  Population Mortality  trends  . . . .  rates  6 6  Production  :  General production  7  i n d i f f e r e n t years  7  P r o d u c t i o n o f c h i c k s i n r e l a t i o n t o age of hen Seasonal d i f f e r e n c e s i n weights o f grouse on b o t h a r e a s  9 11  Movements o f r e s i d e n t m a l e g r o u s e on s p r i n g range . .  12  Behaviour o f t e r r i t o r i a l  cocks i n a t e s t arena  . 13  Grouse a c t i v i t y as r e c o r d e d a t l i s t e n i n g posts:  " t h e s p r i n g o f 1971, 1972, and 1973" . . .  DISCUSSION Population  15 17  trend  i n t h e new p o p u l a t i o n  17  iii TABLE OF CONTENTS  (CONTINUED)  Page  Were t h e new b i r d s b e h a v i o u r a l l y than the o l dstock? Where d o e s t h i s  leave  different 18  us i n e x p l a i n i n g  r e g u l a t i o n o f a n i m a l numbers?  20  SUMMARY  22  LITERATURE CITED  24  iv  L I S T OF TABLES Table 1  Page Density of t e r r i t o r i a l  males  (number/100 a c r e s )  6  2  M o r t a l i t y r a t e s of males  3  Numbers a n d c o m p a r i s o n o f g r o u s e i n h u n t e r s ' b a g s (1969-72) P r o d u c t i o n o f y o u n g on e x p e r i m e n t a l and c o n t r o l a r e a s 1 9 7 1 , 1 9 7 2 , a n d 1973 c o m b i n e d  4  4 (a) 4 (b) 5  (1971-73)  7 8 . 10  Hens w i t h b r o o d s P e r c e n t a g e b r o o d l e s s hens  10 10  Age o f hen and p r o d u c t i o n o f c h i c k s , 1972 and 1973  11  6  R e s p o n s e o f m a l e s t o a r e n a t e s t and c o m p a r i s o n w i t h d a t a o f M o s s o p (1969)  7  A v e r a g e number o f h o o t i n g m a l e s h e a r d per e v e n i n g  . . . .  14 16  1  INTRODUCTION  P r e v i o u s r e m o v a l e x p e r i m e n t s on r e d g r o u s e  (Watson a n d  J e n k i n s , 1968) d e m o n s t r a t e d r a t e s o f r e c r u i t m e n t t h a t  differed  w i t h seasons o f t h e year i n which t h e removal took p l a c e . the  If  r e m o v a l t a k e s p l a c e i n t h e autumn t h e numbers o f b i r d s  occupying t e r r i t o r i e s i n the following spring usually rose t o the  pre-existing density.  The r e d g r o u s e r e m o v a l  experiments  showed a l s o t h a t t h e s i z e o f b r e e d i n g s t o c k s was d e t e r m i n e d by t e r r i t o r i a l  behaviour.  However, no s t u d y s o f a r h a s m o n i t o r e d  a recruitment population o f grouse, a f t e r i n i t i a l to  i n s p e c t f o r emerging changes i n s t o c k q u a l i t y .  recolonization, I t has been  shown t h a t i n a l l g r o u s e - r e m o v a l . e x p e r i m e n t s r e p l a c e m e n t are  drawn f r o m t h e p o o l o f n o n - t e r r i t o r i a l y o u n g In  birds.  1970 B e n d e l l e t a l p e r f o r m e d a r e m o v a l e x p e r i m e n t i n  a population of blue grouse. to  birds  I n 1971 t h e p o p u l a t i o n r e t u r n e d  t h e same d e n s i t y as b e f o r e a n d i n a p p r o x i m a t e l y e v e n s e x  ratio.  A p p r o x i m a t e l y 90% o f t h e r e c r u i t i n g b i r d s w e r e  grouse.  yearling  The r e s u l t s s u b s t a n t i a t e d t h e c o n c l u s i o n t h a t b l u e t  grouse a r e spacing themselves out through t e r r i t o r i a l  behaviour.  The p u r p o s e o f my s t u d y i s t o a n a l y z e t h e q u a l i t y o f r e c r u i t s in  t h e new p o p u l a t i o n . The p r i n c i p a l q u e s t i o n s I a s k e d w e r e : 1. What t r e n d s d i d t h e new p o p u l a t i o n follow? 2.  Were b i r d s e n t e r i n g t h e r e m o v a l a r e a d i f f e r e n t from those e n t e r i n g t h e c o n t r o l area?  2 3.  Where d o e s t h i s l e a v e us i n e x p l a i n i n g t h e r e g u l a t i o n o f a n i m a l numbers?  My s t u d y was c o n d u c t e d on t h e e a s t s i d e o f V a n c o u v e r I s l a n d on t h e same p o p u l a t i o n as r e p o r t e d on by B e n d e l l e t a l ( 1 9 7 2 ) .  3  MATERIALS AND  This of  s t u d y was  1971,  1972,  15 m i l e s  and  c a r r i e d out 1973  west of the  19 70  i n the  springs  town o f C a m p b e l l R i v e r , feet.  I t was  control  on  the  ( M Q L - C ) a r e a s , a p p r o x i m a t e l y one  Removal mile  because of t h e i r  similarity  used to l o c a t e t e r r i t o r i a l o t h e r b i r d s on  the  Population  The  two  apart,  density  resightings  even i n b l u e grouse territorial  be  the  sex  d e t e r m i n e d i n any  m a l e s can  be  the  of hooting  and  presence  of  males per and  one  age year.  100  ratio  Since the  The and  (Bendell,  1967).  acres. and  territorial  banded b i r d s  ( B e n d e l l , 1967), d e n s i t y  c a l c u l a t e d simply  years  been l o g g e d  to detect  of p r e v i o u s l y  f a i t h f u l n e s s with which they r e t u r n successive  were  P o i n t i n g dogs were  f r e q u e n t s i g h t i n g s of newly banded b i r d s .  r e s i g h t data,  and  analysis  m a l e s w e r e b a s e d on  as h o o t i n g  (MQL-X)  areas.  Estimates of population  is  analysis  a r e a s were chosen  i n appearance. m a l e s and  altitude  been removed i n  e s t a b l i s h e d w i t h i n a 4 0 , 0 0 0 - a c r e a r e a t h a t had i n 1952.  a t an  d i r e c t e d t o the  adjacent c o n t r o l area.  l a t e r b u r n e d by w i l d f i r e  summers  approximately  B.C.,  a r e a f r o m w h i c h a l l r e s i d e n t b i r d s had  and  and  a t M i d d l e Quinsam Lake,  o f a p p r o x i m a t e l y 1,000 of the  METHODS  sex  From c a p t u r e  mortality rate of  adult  because of  therefore  be  and  can  to t h e i r t e r r i t o r i e s I t can  ratio  i s expressed  mortality rates  accurately  and  the in  assumed  4  t h a t f a i l u r e to r e c o r d a banded a d u l t male on h i s t e r r i t o r y of  the previous year means t h a t he i s dead.  For a more  d e t a i l e d d e s c r i p t i o n of g e n e r a l methods and see B e n d e l l Stirling  (1955),  Zwickel  (1965),  and B e n d e l l (1966).  (over two  field  procedures  Zwickel and B e n d e l l  (1967),  Grouse were c l a s s i f i e d as a d u l t s  years) or y e a r l i n g s , a c c o r d i n g to the roundness of  the outer two  primaries  c l a s s i f i e d as t e r r i t o r i a l being one  (Boag, 1965).  Male grouse were  or t r a n s i e n t .  f r e q u e n t l y seen i n one  A territorial  locality,  bird  a t r a n s i e n t being  one r e s i g h t e d once or twice over a very l a r g e area, or not being seen again a f t e r banding. y e a r l i n g males and  Body weights of a d u l t and  females were obtained at time of  The weights of y e a r l i n g and a d u l t males and females  banding. on  both  areas were t e s t e d by means of a two-way a n a l y s i s of v a r i a n c e . Behaviour I e v a l u a t e d the q u a l i t y of i n d i v i d u a l s i n two ways: d i r e c t l y by u s i n g t e r r i t o r i a l  movements on the summer range  and the m i r r o r - t e s t i n g of t e r r i t o r i a l  cocks, and  by r e c o r d i n g u n d i s t u r b e d s e x u a l and t e r r i t o r i a l the  indirectly activity in  evenings. a.  Mirror test  behaviour  M i r r o r image s t i m u l a t i o n i s a technique  for studying  a g g r e s s i v e and s o c i a l p a t t e r n s i n a wide v a r i e t y of which respond  to t h e i r r e f l e c t i o n i n a m i r r o r .  on marmots-} f i n c h e s , and parakeets of  animals  animals  Previous s t u d i e s  demonstrated t h a t  responses  to t h e i r r e f l e c t e d image are c o n s i s t e n t w i t h  their  5  behavioural d i s p l a y s noted during s o c i a l i n t e r a c t i o n s i n the field  ( G a l l u p and C a p p e r ,  Mossop  1970;  S v e n d s e n and A r m i t a g e ,  1973).  (1971) u s e d t h e m i r r o r image t e c h n i q u e on t h r e e  p o p u l a t i o n s of b l u e grouse  and e s t a b l i s h e d an i n v e r s e c o r r e l a t i o n  between d e n s i t y o f b l u e grouse m i r r o r images.  different  and r e a c t i o n s t o w a r d s  their  I a p p l i e d t h i s m e t h o d i n t h e s p r i n g o f 1973  examine f o r d i f f e r e n c e s . i n b e h a v i o u r o f t h e t e r r i t o r i a l o f t h e two by  stocks.  males  I p l a c e d t h r e e p l a t e g l a s s m i r r o r s 40  cm  60 cm w e l l w i t h i n t h e t e r r i t o r y o f a m a l e , t o g e t h e r w i t h a  dummy hen b l u e g r o u s e two  f e e t away.  i n squatting copulation position  E a c h t e s t began;.when a m a l e a r r i v e d  to a tape c o n t a i n i n g p r e c o p u l a t o r y c a l l s grouse.  One  t e r r i t o r i a l m a l e was  f r o m a hen  T i m e s f o r t e s t i n g w e r e as f o l l o w s : 2215  about  i n response blue  t e s t e d on t h e e x p e r i m e n t a l  a r e a , a n o t h e r on t h e c o n t r o l a r e a e a c h day  throughout  0600-1030 h o u r s  spring.  and  1930-  hours. b.  G r o u s e b e h a v i o u r r e c o r d e d by o b s e r v e r s i n t h e  L i s t e n i n g p o s t s , from which o b s e r v e r s would o f grouse o n e - h a l f hour p i c k e d on b o t h a r e a s . and summers o f 1972 was  to  and  evenings  record  activity  a f t e r s u n s e t f o r 40 t o 50 m i n u t e s The  same p o s t s w e r e u s e d d u r i n g t h e s p r i n g s  1973,  w h e r e a s i n 1971  behavioural activity  r e c o r d e d w h i l e I w a l k e d on t h e r o a d s i n t h e e v e n i n g s .  intervals,  c a l l s / m i n u t e were r e c o r d e d f o r t h e f i r s t t h r e e  of l i s t e n i n g . grouse  calls  D u r i n g the r e m a i n i n g seven minutes,  At  10-min  minutes  intensity  of  and t h e numbers o f g r o u s e h e a r d w e r e n o t e d .  Listening started early i n April o f -June.  were  and c o n t i n u e d u n t i l t h e  middle  6  RESULTS  Population  trends  B e t w e e n 1970 a n d 1973 g r o u s e  numbers i n t h e C a m p b e l l  R i v e r area dropped from s i x t o f o u r t e r r i t o r i a l 100  acres  (Table 1 and B e n d e l l ,  TABLE 1.  males p e r  1972).  D e n s i t y o f t e r r i t o r i a l m a l e s (no./lOO a c r e s ) . A l l b i r d s on t h e e x p e r i m e n t a l a r e a (MQL-X) w e r e removed i n 1970. MQL-X  No.  (90 0 a c r e s )  of Birds  MQL-C  Density  No.  (800 a c r e s )  of Birds  Density  1970  64  6.1  48  6.0*  1971  46  5.1  52  6.5  1972  38  4.1  45  5.6  1973  35  4.0  30  3.8  This t o t a l  Mortality  i sprobably  rates The  mortality rate of adult t e r r i t o r i a l  was  approximately  for  other blue grouse  in  l o w , a s t h e c e n s u s was i n c o m p l e t e  30% p e r y e a r , w h i c h populations  agrees  o n l y i n 1971,  ( T a b l e 2)  with that reported  ( B e n d e l l , 19 67) .  t h e p o p u l a t i o n appears t o r e s u l t from  y e a r l i n g s being inadequate  males  The d e c l i n e  the recruitment of  to replace the losses of adults:  on t h e c o n t r o l a r e a d i d e n o u g h y e a r l i n g s e n t e r  7  t h e p o p u l a t i o n t o p r e v e n t numbers  TABLE 2.  from  M o r t a l i t y r a t e s o f males  dropping.  (1971-73)  MQL-X  MQL-C  No. Banded  No. Returned  % Mortality  No. Banded  No. Returned  % Mortality  territorial adults  22  16  28  53  41  23  territorial yearlings  16  15  8  7  6  14  Production T h e r e i s much v a r i a t i o n and  from year t o year i n t h e p r o d u c t i o n  s u r v i v a l o f y o u n g t o autumn, b u t p a s t s t u d i e s o f b l u e  grouse  h a v e a l l shown t h a t t h e r e a r e more t h a n enough  the f o l l o w i n g s p r i n g t o r e p l a c e a d u l t m o r t a l i t y 1955;  Redfield,  Bendell,  yearlings  (Bendell,  1972; Z w i c k e l a n d B e n d e l l , 1967; Z w i c k e l a n d  1972).  General production i n d i f f e r e n t  years  P r o d u c t i o n o r b r e e d i n g s u c c e s s was d e f i n e d f o r t h e p u r p o s e s of  t h i s s t u d y a s number o f c h i c k s r a i s e d p e r b r o o d b y A u g u s t .  In  1971, b r o o d s  were t h e s m a l l e s t o f t h e f o u r y e a r s  (1970-1973).  However, when I c o m p a r e d p r o d u c t i o n d a t a f o r 1971 w i t h t h e p r o d u c t i o n d a t a f o r 1970, 1972, a n d 1973, I f o u n d no s i g n i f i c a n t d i f f e r e n c e i n numbers o f b r o o d s  observed,  c h i c k s p e r brood e a r l y i n t h e season.  o r i n numbers o f  T h e r e was a l s o no  8  s i g n i f i c a n t d i f f e r e n c e i n t h e p r o p o r t i o n o f c h i c k s found i n t h e h u n t e r road check d a t a comparing and 1 9 7 2 a g a i n s t 1 9 7 1 (Table 3 ) .  in1969, 1970,  hunter k i l l  The percentage o f j u v e n i l e s  shot p e r a d u l t o r y e a r l i n g female was s i g n i f i c a n t l y l o w e r . Another measure o f p r o d u c t i o n i s t h e p e r c e n t a g e o f b r o o d l e s s hens observed l a t e r i n t h e season August) a f t e r most broods have h a t c h e d . may  r e s u l t from:  the i n a b i l i t y  ( i ; e . J u l y and  A b r o o d l e s s hen  i n f e r t i l e eggs, l o s s o f young, n e s t p r e d a t i o n ,  t o r e n e s t , n e s t d e s e r t i o n , and t h e p h y s i o l o g i c a l  i n a b i l i t y t o produce young.  The number o f b r o o d l e s s hens  observed i n J u l y and August i n 1 9 7 1 on b o t h areas was s i g n i f i c a n t l y h i g h e r than i n t h e o t h e r t h r e e y e a r s  TABLE 3 .  Numbers and comparison bags ( 1 9 6 9 - 1 9 7 2 )  (Table 4 ( b )  P / . O O I ) .  o f grouse i n h u n t e r s '  1969  1970  1971  1972  54  57  22  45  Number o f h u n t e r s  39  38  58  40  % chicks  68  69  40  73  Total birds  tallied  The removal birds.  a r e a (MQL-X) i n  1971  had o n l y y e a r l i n g  S i n c e y e a r l i n g cocks a r e though t o be  less f e r t i l e  and y e a r l i n g hens p o o r e r mothers because o f i n e x p e r i n c e o r physiological inability expected  (Bendell,  t o produce,  1967).  The y e a r  fewer broods might be 1971  p r o d u c t i o n o f c h i c k grouse on b o t h a r e a s .  r e s u l t e d i n poorer On MQL-X however,  9  4 3 hens p r o d u c e d t h r e e b r o o d s o n l y  ( 7 % s u c c e s s ) w h i l e on t h e  c o n t r o l a r e a 32 f e m a l e s p r o d u c e d e i g h t b r o o d s  (25% s u c c e s s )  ( T a b l e 4 [Ja] ) . T h i s d i f f e r e n c e i s s t a t i s t i c a l l y The  two a r e a s w e r e c h o s e n b e c a u s e  significant.  t h e y were s i m i l a r ,  and  t h e r e i s no e v i d e n c e t h a t e n v i r o n m e n t a l d i f f e r e n c e s w e r e a s s o c i a t e d w i t h the d i f f e r e n c e s , i n  success.  I therefore  c o n c l u d e t h a t t h e p o o r s u c c e s s on t h e e x p e r i m e n t a l a r e a i n 1971 was  due  t o t h e hens a l l b e i n g y e a r l i n g s .  Production of chicks i n r e l a t i o n The of  t o age o f  s e a r c h f o r b r o o d s i n 19 73 was  hen  t e r m i n a t e d by t h e e n d  J u n e , t h e r e f o r e t h e numbers o f b r o o d s f o u n d i n 1972  1973 w e r e added  t o g e t h e r t o i n c r e a s e t h e sample  size.  and  early  The  hens  on t h e r e m o v a l a r e a p r o d u c e d p r o p o r t i o n a t e l y more b r o o d s and more c h i c k s p e r b r o o d  (2.6) i n 1972  and 1973  than d i d the  h e n s on t h e c o n t r o l a r e a , w h i c h p r o d u c e d o n l y 16 b r o o d s an a v e r a g e o f 1.7' c h i c k s p e r b r o o d  (Table 5).  (27)  Because  with of the  r a p i d r e c o l o n i z a t i o n i n 1 9 7 1 , t h e b r e e d i n g h e n s on MQL-X i n 1972  and 1973 w e r e p r i m a r i l y  lings. of  2 - y e a r - o l d h e n s and a few y e a r -  T a b l e 5 shows t h e p e r c e n t o f b r o o d s p r o d u c e d by h e n s  d i f f e r e n t ages.  The  2 - y e a r - o l d hens, the m a j o r i t y  w h i c h were t h e o r i g i n a l r e c r u i t s  of  i n 1971, a c c o u n t e d f o r t h e  g r e a t e r number o f b r o o d s and t h e g r e a t e r number o f  chicks  per  revealed  b r o o d on t h e r e m o v a l a r e a .  F i e l d observations  no d i f f e r e n c e i n t h e p r o p o r t i o n o f h e n s t h a t d e f e n d e d b r o o d s , or the  i n the behaviour of the c h i c k s .  We  conclude therefore,  d i f f e r e n c e i n b r e e d i n g s u c c e s s i s due n o t t o t h e  that  differences  10  P r o d u c t i o n o f y o u n g on e x p e r i m e n t a l and c o n t r o l a r e a s 1971, 1972, a n d 1973 c o m b i n e d  TABLE 4.  TABLE 4(a)  Hens w i t h  brood Control  Experimental > 10 d a y s o l d c/b* 1971  (N)<10 days o l d c/b  3.0  (3)**  0  3.0  (9)  3.3  (N)  1. 25  (18)  1.5  (4)  2. 0  c/b d e n o t e s c h i c k s p e r b r o o d Number o f i n d i v i d u a l b r o o d s  Percentage broodless  1971 1972 & 1973  removal  (12)  hens  Experimental  1970  (7)  (8)  * **  Broodless  (N)  2.8  &  With brood  (N) <10 d a y s o l d c/b  (0)  1972 1973  TABLE 4 (b) .  >10 d a y s old'c/b  Control  % Broodless  year  With Broodless brood  % Broodless  39  22  36  3  39  92  8  24  75  18  2  10  11  4  26  11  i n q u a l i t y b u t i n age s t r u c t u r e o f b r e e d i n g h e n s  TABLE 5.  Age o f h e n a n d p r o d u c t i o n o f c h i c k s , Removal  Age o f A v e r a g e hen no. o f (years) c h i c k s  No. o f broods  ( T a b l e 5)  1972 a n d 1973  Control % of total broods  Average no. o f chicks  11  0  63  1.4  7  44  No. o f broods  % of total broods  1**  3.6  2  3.1  17  3  2.6  5  18.5  1.5  2  12  2  7.5  2.3  7  44  > 2 (Bird?-1.5 b a n d e d as adult) Total  **  2.6  ~2A  0  1.7  1 y e a r o l d hens o m i t t e d from comparison c h i c k s w i t h age o f h e n .  16  of production of  S e a s o n a l d i f f e r e n c e s i n w e i g h t s o f g r o u s e on b o t h  areas  S e a s o n a l t r e n d s i n w e i g h t w e r e t h e same f o r b o t h MQL-X and MQL-C.  R e g a r d l e s s o f s e x o r age,  and a l l e x h i b i t e d a s u b s e q u e n t summer, as r e p o r t e d b y R e d f i e l d of  a l l showed a p e a k i n May  d e c l i n e toward t h e end o f (1972).  I n g e n e r a l , no t r e n d  s i g n i f i c a n c e was f o u n d i n a n y age o r s e x c a t e g o r y .  There  w e r e no d i f f e r e n c e s i n mean w e i g h t s b e t w e e n a r e a s i n t h e y e a r s 1970,  1 9 7 1 , 1 9 7 2 , a n d 1973.  on t h e r e m o v a l  Therefore, I.conclude that  a r e a w e r e n o t u n d e r a n y more s t r e s s ,  by w e i g h t s , t h a n t h o s e f o u n d o n t h e c o n t r o l .  grouse  as j u d g e d  12  Movements o f r e s i d e n t male grouse on s p r i n g range B l u e grouse a r e m i g r a t o r y .  However, once on t h e s p r i n g  range t h e r e s i d e n t b r e e d i n g males occupy t e r r i t o r i e s t o w h i c h they.return year a f t e r year  (Bendell, 1955).  R e s i d e n t males  move about w i t h i n t h e b o u n d a r i e s o f t h e t e r r i t o r y t o f e e d , b r e e d , and p e r f o r m o t h e r a c t i v i t i e s such a h o o t i n g , and e x c l u d i n g o t h e r male grouse.  Through s y s t e m a t i c  fighting, search,  banded r e s i d e n t s were l o c a t e d p e r i o d i c a l l y and t h e i r l o c a t i o n s p l o t t e d on a l a r g e f i e l d map.  The s i z e o f an o c c u p i e d t e r r i t o r y  i s d e t e r m i n e d by r e p e a t e d s i g h t i n g s o f i n d i v i d u a l l y males.  identifiable  I n s p r i n g , a d u l t males a r e v o c a l and t e r r i t o r i a l  most y e a r l i n g males a r e s i l e n t and move about w i d e l y  while  (Bendell  and E l l i o t , 1 9 6 7 ) , hence most y e a r l i n g s a r e c l a s s i f i e d as t r a n s i e n t s unless  they e x h i b i t s t r o n g t e r r i t o r i a l  tendencies  and a r e r e s i g h t e d f r e q u e n t l y i n t h e same l o c a t i o n . In 1971, 1970,  a f t e r the k i l l i n g of a l l t e r r i t o r i a l b i r d s i n  t h e removal a r e a c o n t a i n e d  ( B e n d e l l £t a l , 1 9 7 2 ) .  a p p r o x i m a t e l y 90% y e a r l i n g s  The y e a r l i n g cocks wandered i n a s i m i l a r  manner t o t h a t o f y e a r l i n g cocks i n a r e s i d e n t c o n t r o l of mixed ages, o r t h e r e s i g h t i n g d i s t a n c e s b e i n g e v e n l y b u t e d w i t h i n t h e range o f 20 t o 2,000 y a r d s I n 1972,  population distri-  (unpublished  data).  t h e i r f i r s t y e a r i n a d u l t plumage, t h e b i r d s became  territorial  a f t e r t h i s placement and i n 1973,  as t h e age  s t r u c t u r e o f t h e removal a r e a approached t h a t o f t h e c o n t r o l , the a d u l t t e r r i t o r i a l males on b o t h areas were comparable i n the d i s t a n c e t r a v e l l e d between r e s i g h t i n g s , the m a j o r i t y  being  r e l i g h t e d w i t h i n 20 t o 70 y a r d s o f where they were i n i t i a l l y  13  recorded. Males which had r e c o l o n i z e d t h e removal t h e r e s i d e n t males on t h e c o n t r o l a r e a .  a r e a behaved  like  A l t h o u g h i n 1971  MQL-X was v a c a n t , y e a r l i n g s t h a t moved i n b e h a v e d as n o r m a l yearlings, showing  some b e i n g n o m a d i c i n t h e i r movements a n d a f e w  territorial  were t e r r i t o r i a l  behaviour.  on t h e r e m o v a l  When t h e same b i r d s ,  as a d u l t s ,  a r e a i n 1972 a n d 1973 t h e y  o c c u p i e d a p p r o x i m a t e l y t h e same s i z e t e r r i t o r i e s  as t h e b i r d s  on t h e c o n t r o l . Behaviour o f t e r r i t o r i a l Three  cocks i n a t e s t  arena  main r e a c t i o n s were r e c o g n i z e d i n t h e t e s t  "exploratory",  " c o u r t i n g " , and " i n t e r a c t i o n " .  arena:  Exploratory  b e h a v i o u r was t h e r e a c t i o n o f a m a l e t o t h e w h i n n y c a l l o f a female blue grouse  ( S t i r l i n g and B e n d e l l , 1970).  T h e r e was  e i t h e r no r e s p o n s e , e v i d e n c e d b y no movement o r a l t e r a t i o n i n his  h o o t i n g r a t e , o r a r e s p o n s e w h i c h was c h a r a c t e r i z e d b y  an a p p r o a c h  and a n o t i c e a b l e change i n h o o t i n g r a t e  (Table 6 ) .  T h e r e was no s i g n i f i c a n t d i f f e r e n c e i n t h e p e r c e n t a g e o f h o o t i n g territorial  males t h a t f a i l e d t o respond t o t h e t e s t  on t h e c o n t r o l a n d r e m o v a l The  approach  areas  ( T a b l e 6, P = . 2 6 ) .  b y a m a l e t o t h e t e s t a r e n a was b y w a l k i n g ,  running, or f l y i n g ,  and was u s u a l l y a c c o m p a n i e d b y a s l i g h t  i n c r e a s e i n t h e number o f h o o t i n g p h r a s e s which  t h e y w e r e made.  was n o t s i g n i f i c a n t .  for  o f the males,  between  but the difference  A p p r o x i m a t e l y 80% o f t h e males on b o t h  o r r a n i n t o the t e s t arena.  r a t e o f approach  and t h e i n t e n s i t y w i t h  T h e r e was a s l i g h t d i f f e r e n c e  the areas i n t h e r a t e o f approach  areas walked  situation  was s i g n i f i c a n t l y  Mossop's  (1971)  figure  g r e a t e r than the f i g u r e s  I calculated. the  However, he d i d n o t s t a t e how t h e b i r d s  arena. Analysis  o f t h e c o u r t i n g phase i n v o l v e d  number o f f e m a l e - o r i e n t e d  behaviours during  i n g them a s a c t s o b s e r v e d p e r m i n u t e . was by  t h e b i r d s o n t h e two a r e a s  to that  t h e t e s t and e x p r e s s -  No s i g n i f i c a n t d i f f e r e n c e exhibited  i n t e r a c t i o n p h a s e was c o m p u t e d i n a s i m i l a r manner f o r t h e c o u r t i n g phase, a l l aggressive  acts p e r minute. aggression rates ically  acts  ( P ^ 0.07) .  towards t h e r e f l e c t i o n i n t h e m i r r o r  (P< 0 . 0 0 2 ) .  TABLE 6.  directed  being expressed  Although t h i s  finding.  ( T a b l e 6) i n d i c a t e no d i f f e r e n c e i n t h e two  as lower  figure i s statist-  additional tests are required  biological significance of this  males l i v i n g  acts  The r e m o v a l a r e a h a d s i g n i f i c a n t l y  significant,  these t e s t s the  counting the  observed i n average frequency o f c o u r t i n g  The  the  approached  t o reveal  Results  ofa l l  i n behaviour of  populations.  Response o f males t o a r e n a t e s t and comparison d a t a o f Mossop (1969) MQL-X  MQL-C  MQL-X Mossop  with  (1969)  % response  71 (21)  76 (17)  average rate of advance  12.3 (9) SD = 5.00  9.0 (7) SD = .6.38  21.73 (16) SD = 4.38  average aggressive acts/min  11.74 (6) SD = 5.96  1 3 . 1 (5) SD = 3.7  11.0 (8)  average rate of courting  5.85 (4) SD = 0.57  5.20 (4) SD = 0.23  5.76 (7) SD = 2.60  15  Grouse a c t i v i t y 1971,  1972,  as r e c o r d e d a t l i s t e n i n g  and  posts' (springs of  1973)  L i s t e n i n g a t r e g u l a r p o s t s gave a measure o f grouse and In  activity  listening  throughout  from roads  t h e s e a s o n s o f 1972  and  d e c l i n e s i n a v e r a g e numbers o f h o o t i n g m a l e s  and  a v e r a g e numbers o f h o o t s flutters.  per minute, I n 1971,  total  (Table  were t h o u g h t  t o be  less vocal  most y e a r l i n g  incapable of hooting, whooting,  f u l c o p u l a t i o n ( B e n d e l l and E l l i o t ,  s t u d y , h o w e v e r , t h e new hoot  1967).  males and  In the  y e a r l i n g s on t h e r e m o v a l  classified  g i v e an a c c u r a t e a c c o u n t  as w h o o t s and  hen  successpresent  area i n  few i n d i v i d u a l b i r d s w h i c h evenings.  trends throughout  calls  do  o f t h e w h o l e p o p u l a t i o n on an  because the b u l k of these b r e e d i n g a c t i v i t i e s  two  after  1971  regularly.  Activities  or  the  (Table 7 ) .  I n p a s t s t u d i e s on b l u e g r o u s e ,  did  hen  when m o s t m a l e s on  t h a n w e r e t h e m a l e s on t h e c o n t r o l a r e a , e s p e c i a l l y  ful  7)  number o f  a r e a were y e a r l i n g s , t h e y were s l i g h t l y  mid-May  1971.  showed s i m i l a r s e a s o n a l p e a k s  and  removal  1973,  s u p p l i e d a s i m i l a r index i n  a l l three years both areas  c a l l s , w h o o t s , and  evening  not area  involve only  h a p p e n e d t o be v e r y a c t i v e on  one  Both p o p u l a t i o n s were s i m i l a r i n h o o t i n g 1972  and  19 73  (Table 7 ) .  a  TABLE 7.  A v e r a g e number o f h o o t i n g m a l e s h e a r d p e r e v e n i n g (40 t o 50 m i n u t e s e v e r y d a y , o n e - h a l f h o u r a f t e r sunset) Experimental  Control  1971  1972  1973  1971  1972  April  1.3  1.0  2.0  1.3  1.5  2.0  May  1.6  2.5  2.1  2.2  2.75  3.25  June  0.7  1.6  1.7  2.1  2.0  1.6  1973  17  DISCUSSION  Previous  s t u d i e s on b l u e g r o u s e ,  parasites, predators,  f o o d , and  f o c u s i n g on  v e g i t a t i o n have f a i l e d t o e x p l a i n  w h a t d e t e r m i n e s t h e numbers o f g r o u s e l i v i n g dell,  1967;  1972).  B e n d e l l and  Studying  approach to the study authors  Elliot,  the behaviour  1967;  and  of animals  i n an a r e a .  Z w i c k e l and  of population r e g u l a t i o n .  K i n g , 1968;  Social interaction, behaviour,  Sadleir,  1965;  e x h i b i t e d through  Bendell,  which  (Healey,  Watson,  t h e i r breeding  For t h i s behaviour  range  are  1967;  1965).  possible agonistic  t a k e s p l a c e i n b l u e g r o u s e i n t h e s p r i n g when  a p p e a r on  new  A number o f  have r e c e n t l y r e p o r t e d changes i n b e h a v i o u r  J e n k i n s , 1961;  (Ben-  is a relatively  c o r r e l a t e d w i t h changes i n p o p u l a t i o n d e n s i t y  first  diseases,  ( B e n d e l l and  they  Elliot,  1967).  to regulate a population i t s expression  must be d e n s i t y - r e l a t e d . P r e v i o u s b l u e and that i n every  red grouse removal experiments i n d i c a t e d  i n s t a n c e , regardless of area or d e n s i t y of  s t o c k , c o m p l e t e r e c r u i t m e n t a l w a y s o c c u r r e d by spring. 1972).  ( B e n d e l l e t a l 1972; However, no  W a t s o n and  the f o l l o w i n g  J e n k i n s , 1968;  f o l l o w - u p s t u d i e s have m o n i t o r e d  P o p u l a t i o n t r e n d i n t h e new  Zwickel,  the  r e c r u i t m e n t p o p u l a t i o n f o r l a t e n t changes i n p o p u l a t i o n o r d i f f e r e n c e s i n t h e new  breeding  trends  stock. population  A general d e c l i n i n g t r e n d of grouse p o p u l a t i o n s  i n the  18  Campbell  R i v e r a r e a was r e v e a l e d t h r o u g h a l o n g - t e r m  p r i o r t o the removal experiment.  study  I n 1971, t h e r e c o l o n i z i n g  p o p u l a t i o n e s t a b l i s h e d t h e same d e n s i t y as t h e o l d p o p u l a t i o n , and  i n 1972 a n d 1973 c o n t i n u e d t o d e c l i n e , p a r a l l e l i n g t h e  trend i n the surrounding control area.  The a p p e a r a n c e  p e r s i s t e n c e o f t h e s e new b i r d s on t h e r e m o v a l  and  area d i dnot  r e s u l t i n a l o w e r r e c r u i t m e n t r a t e on s u r r o u n d i n g a r e a s , hence t h i s  s u p p o r t e d t h e c o n c l u s i o n t h a t t h e s e new  recruits  w e r e s u r p l u s ( B e n d e l l e t a l , 1 9 7 2 ) , and t h a t t h e g e n e r a l decline d i d not a r i s e of p o t e n t i a l Three experiment  f r o m t h e r e b e i n g an i n s u f f i c i e n t number  recruits.  outcomes might  have been e x p e c t e d from t h e removal  i fbehaviour i s involved i n the l i m i t a t i o n of density  i n an e s t a b l i s h e d g r o u s e p o p u l a t i o n .  Replacement  individuals  c o u l d be more o r l e s s a g g r e s s i v e t h a n t h o s e r e m o v e d , i n w h i c h case they might  s e t t l e more o r l e s s c l o s e l y  change a c c o r d i n g l y . ness and s e t t l e experiment  T h i r d l y , t h e y c o u l d be o f s i m i l a r ,  a t t h e same d e n s i t y .  revealed.  and numbers  would aggressive-  T h i s i s i n d e e d what t h e  These d a t a a r e e v i d e n c e a g a i n s t a b e h a v i o u r -  a l d i f f e r e n c e i n the replacement  stock.  Were t h e new b i r d s b e h a v i o u r a l l y d i f f e r e n t f r o m t h e o l d s t o c k ? A g g r e s s i o n , one f a c t o f a g o n i s t i c b e h a v i o u r , i s known t o affect l i t t e r  s u r v i v a l i n mice  ( S o u t h w i c k , 1955) a n d t o be  c o r r e l a t e d w i t h changes i n p o p u l a t i o n d e n s i t y i n r e d grouse (Watson and M o s s , 1 9 6 9 ) , b l u e g r o u s e mice  (Krebs, 1970).  (Mossop, 1 9 7 1 ) , a n d  The r e c o l o n i z i n g g r o u s e o n t h e e x p e r i -  me-ntal a r e a h a d s i m i l a r  s u r v i v a l r a t e s and w e i g h t s  when  when c o m p a r e d w i t h t h e g r o u s e on t h e c o n t r o l a r e a . from l i s t e n i n g p o s t s and a r e n a  Results  t e s t i n g y i e l d e d no e v i d e n c e o f  b e h a v i o u r a l d i f f e r e n c e s between a r e a s .  I f t h e r e had been any  o u t s t a n d i n g d i f f e r e n c e s between p o p u l a t i o n s , they  would  h a v e e m e r g e d a s a g e n e r a l t r e n d i n t h e p o p u l a t i o n i n 19 72 and  1973.  Furthermore,  d i f f e r e n t l y behaved i n d i v i d u a l s  would  h a v e r e c o l o n i z e d t h e a r e a a t a d i f f e r e n t d e n s i t y o r i n some way t y p i f i e d  a d i f f e r e n t l y behaved p o p u l a t i o n .  One o f t h e d i f f e r e n c e s f o u n d  between areas  concerned  the  w i l d n e s s o f c e r t a i n b i r d s on t h e r e m o v a l a r e a i n 1971 and 1972  such  t h a t i t was d i f f i c u l t  tion  from these  individuals  to obtain behavioural  i n the f i e l d  or i n the test  However, a n a l y s i s o f u n d i s t u r b e d a c t i v i t y posts near these higher o r lower those  informaarena.  at the listening  i n d i v i d u a l s d i d n o t e x p o s e any r e c o g n i z a b l y evening  activity  i n the control population.  r a t e s when c o m p a r e d w i t h I f these uncatchable  birds  represent a p o r t i o n of the population that i s g e n e t i c a l l y d i f f e r e n t t h e r e s h o u l d be a d i f f e r e n c e i n s e t t l i n g p a t t e r n o r s i g h t i n g s i n t h e a r e a s o c c u p i e d by t h e s e c l u s i v e b i r d s . T h i s h o w e v e r was n o t o b s e r v e d .  Therefore,  the presence  of  c e r t a i n w i l d b i r d s i n g r o u s e p o p u l a t i o n s may be t h e r e s u l t of repeated learning to  unsuccessful capture  to aviod the observers.  human o b s e r v e r s ;  other  attempts  and o f t h e grouse  The b i r d s a p p e a r e d  wild  they d i d n o t n e c e s s a r i l y appear w i l d t o  grouse. The  o t h e r d i f f e r e n c e i n t h e r e c r u i t m e n t p o p u l a t i o n was  20 the  p r o d u c t i o n o f c h i c k s on the areas i n 1972 and 1973.  The  new p o p u l a t i o n produced what appeared t o be more broods and c h i c k s than the c o n t r o l p o p u l a t i o n , but a l l f i e l d o b s e r v e r s r e c o r d e d no v i s i b l e d i f f e r e n c e i n b e h a v i o u r o f hens g i v i n g brood-defense o r h e n - c h i c k i n t e r a c t i o n when d i s t u r b e d i n the field.  In 19 71,  Mossop found t h a t areas c o n t a i n i n g h i g h  d e n s i t i e s o f grouse a l s o had h i g h s u c c e s s i n p r o d u c t i o n o f chicks.  W i t h no d i f f e r e n c e i n b e h a v i o u r . o f hens o r c h i c k s  e v i d e n t between t h e removal and c o n t r o l a r e a s the b e t t e r p r o d u c t i o n o b s e r v e d on the removal a r e a , a l t h o u g h n o t s t a t i s t ically the  s i g n i f i c a n t , i s a t t r i b u t e d t o the age s t r u c t u r e o f  b r e e d i n g hens.  Where does t h i s l e a v e us i n e x p l a i n i n g r e g u l a t i o n o f a n i m a l numbers? The removal e x p e r i m e n t demonstrated t h a t r e s i d e n t b i r d s r e s t r i c t e d new b i r d s from s e t t l i n g on t h e i r t e r r i t o r y o r nearby. The new r e c r u i t s , f r e e d o f s u p p r e s s i o n from e s t a b l i s h e d  birds,  might have s e t t l e d t o a h i g h e r p o p u l a t i o n d e n s i t y than e x i s t e d there before.  I n f a c t , though they wandered more w i d e l y and  showed l e s s tendency t o e s t a b l i s h t e r r i t o r i e s , t h e i r  overall  d e n s i t y d i d not d i f f e r * f r o m t h a t o f the male p o p u l a t i o n removed. The new b i r d s a l s o appeared t o be r e g u l a t e d from- the. y e a r they r e c o l o n i z e d . regulation.  S e t t l i n g i n even s e x - r a t i o i s e v i d e n c e o f  S i n c e b l u e grouse do not form p a i r bonds t h e r e  seems t o be no reason why  i n i t i a l l y t h e r e would have not been  unequal sex r a t i o , t h a t i s assuming t h a t i n any y e a r t h e r e may be more s u r p l u s o f one sex than the o t h e r . known how much i s s u r p l u s nor how  I t i s not  the new p o p u l a t i o n a t t a i n e d  21  sexual e q u a l i t y .  However, i t appears t h a t s e x u a l m a t u r i t y  and the presence  of e x p e r i e n c e d b i r d s are not n e c e s s a r y f o r  a population to regulate i t s e l f . The r e s u l t s o f t h i s study r a t e d the r e c o l o n i z e r s as a g g r e s s i v e and i n no way i n surrounding areas.  b e h a v i o u r a l l y d i f f e r e n t from grouse  The  c h a r a c t e r i s t i c b e h a v i o u r o f the  grouse i n t h a t a r e a has n o t changed s i n c e Mossop (1971)  conducted  m i r r o r t e s t s on the o l d r e s i d e n t s on the removal a r e a i n  1969.  T h e r e f o r e , t h e i n d i v i d u a l s r e c r u i t e d t o the Campbell R i v e r a r e a are b e h a v i o u r a l l y c h a r a c t e r i s t i c of the  surrounding  residents. F u t u r e r e s e a r c h i n t o r e g u l a t i o n o f grouse p o p u l a t i o n s s h o u l d c o n c e n t r a t e on s t u d i e s a t d i f f e r e n t p o p u l a t i o n d e n s i t i e s c o n d u c t i n g m a n i p u l a t i o n experiments of grouse.  o f the h a b i t a t and  stocks  More u n d e r s t a n d i n g o f s o c i a l i n t e r a c t i o n w i t h i n  a p o p u l a t i o n would be a c h i e v e d through e x p e r i m e n t i n g t o f i n d out how  much s u r p l u s i s a v a i l a b l e i n the s p r i n g .  Female  grouse,  because o f t h e i r s e c l u s i v e b e h a v i o u r , are the l e a s t known p o p u l a t i o n component i n the s p r i n g t i m e .  Females are  capable  of d i s p l a y i n g a g g r e s s i o n , p a r t i c u l a r l y p r i o r t o e g g - l a y i n g season ( S t i r l i n g ,  1962;  Theberge, u n p u b l i s h e d Ph.D.  thesis,  1971), and s e v e r a l i n c i d e n t s i n v o l v i n g a g g r e s s i o n amongst females have been r e c o r d e d i n the f i e l d and i n the a v i a r y . Research on females would be v a l u a b l e i n e x p l a i n i n g e q u a l sex r a t i o , the v a r i a b l e compensation f o r m o r t a l i t y w i t h i n the sexes, and i n g e n e r a l would r e s u l t i n a b r o a d e r ~on-"population r e g u l a t i o n .  outlook  22  SUMMARY  The p u r p o s e o f t h i s  s t u d y was  p o p u l a t i o n of blue grouse.  to analyze a r e c o l o n i z i n g  D i f f e r e n t techniques  p o p u l a t i o n census to mirror-image  of r e s i d e n t grouse.  t h e same d e n s i t y as t h o s e even sex r a t i o . declining,  from  s t i m u l a t i o n were used t o t r y  to evaluate the aggressive behaviour the removal  ranging  of the recolonizers  after  The new b i r d s r e c o l o n i z e d t o  i n the surrounding  areas  and i n  The p o p u l a t i o n i n t h e a r e a was g e n e r a l l y  and c o n t i n u e d t o d e c l i n e t h r o u g h o u t  the study.  The a n n u a l m o r t a l i t y r a t e o f a d u l t s was s i m i l a r on t h e c o n t r o l and  experimental areas  (approximately 30%).  In the  first  y e a r t h e e x p e r i m e n t a l a r e a was c o l o n i z e d by 9 0 % y e a r l i n g  birds.  I n o t h e r y e a r s t h e r e c r u i t m e n t r a t e s on t h e e x p e r i m e n t a l and  c o n t r o l a r e a s were l e s s  than  30%.  Production of chicks i n  1972 and 1973 was s l i g h t l y b e t t e r on t h e e x p e r i m e n t a l t h a n on t h e c o n t r o l a r e a , b u t t h i s was n o t significant.  statistically  No d i f f e r e n c e i n l i v e w e i g h t s  on t h e two a r e a s was  found  area  of birds  caught  r e g a r d l e s s o f age o r s e x o f b i r d s  sampled. B e h a v i o u r a l t e s t s on r e s i d e n t m a l e s no d i f f e r e n c e i n r a t e o f a p p r o a c h , between the a r e a s . s i m i l a r t o those  i n an a r e n a  showed  courting, or aggression  R e s u l t s o b t a i n e d from t h e t e s t were  found  i n 1971 by M o s s o p , who  t e s t s on t h e o l d r e s i d e n t p o p u l a t i o n .  conducted  arena  The  g r o u s e t h a t now  essentially  occupy  t h e same t y p e o f g r o u s e  and i n t h e o l d p o p u l a t i o n . limited  the experimental area are  The  new  found i n s u r r o u n d i n g areas p o p u l a t i o n appeared  and r e g u l a t e d f r o m t h e f i r s t y e a r o f  recolonization  d e s p i t e t h e a b s e n c e o f m a t u r e and e x p e r i e n c e d b i r d s .  In  b l u e g r o u s e , r e g u l a t i o n t a k e s p l a c e i n e a r l y s p r i n g on b r e e d i n g range, where d i s p u t e o v e r t e r r i t o r i e s ^ i n v o l v e s r e s i d e n t s and p o t e n t i a l  recruits.  t o be  the both  24  LITERATURE CITED  B e n d e l l , J . F . , 1955. Age, b r e e d i n g b e h a v i o u r and m i g r a t i o n o f s o o t y g r o u s e , Dendragapus o b s c u r u s f u l i g i n o s u s (Ridgway). T r a n s . N. Am. W i l d l . C o n f . , 2 0 : 3 6 7 - 3 8 1 . B e n d e l l , J . F . , a n d P.W. E l l i o t , 1 9 6 7 . B e h a v i o u r a n d r e g u l a t i o n o f numbers i n b l u e g r o u s e . Can. W i l d l . S e r v . Report S e r . No. 4, 76 p p . B e n d e l l , J . F . , D.G. K i n g , a n d D.H. M o s s o p , 19 72. Removal and r e p o p u l a t i o n o f b l u e grouse i n a d e c l i n i n g p o p u l a t i o n . J . W i l d l . Mgmt. 36. No. 4,pp. 1 1 5 3 - 1 1 6 5 . B o a g , D.A., 1 9 6 5 . I n d i c a t o r s o f s e x , age a n d b r e e d i n g p h e n o l o g y i n blue grouse. J . W i l d l . Mgmt. 29:103-108. G a l l u p , G.G., a n d S t u a r t A. C a p p e r , 1 9 7 0 . P r e f e r e n c e f o r m i r r o r image s t i m u l a t i o n i n f i n c h e s a n d p a r a k e e t s . Anim. Behav., 18:621-624. H e a l e y , M.C., 1 9 6 7 . A g g r e s s i o n a n d s e l f - r e g u l a t i o n o f p o p u l a t i o n s i z e i n deermice. E c o l . , 48:377-392. J e n k i n s , D., 1 9 6 3 . P o p u l a t i o n c o n t r o l i n r e d g r o u s e . I n t e r n . O r n i t h . C o n g r . , pp. 690-700.  Proc. 13th  Krebs, C . J . , 1970. B e h a v i o u r a l changes a s s o c i a t e d w i t h t h e population cycle i n voles. E c o l , 5 1 , No. 1. M o s s o p , D. H. 1 9 7 1 . A r e l a t i o n b e t w e e n a g g r e s s i v e b e h a v i o u r and p o p u l a t i o n d y n a m i c s i n b l u e g r o u s e . Unpubl. M.Sc. t h e s i s , U n i v e r s i t y o f B.C., 119 p p . R e d f i e l d , J . A . , F.C. Z w i c k e l , a n d J . F . B e n d e l l , 1 9 7 1 . E f f e c t s o f f i r e on numbers o f b l u e g r o u s e . Proc. T a l l Timbers F i r e E c o l . Conf. 10:63-83. R e d f i e l d , J.A., 1972. V a r i a t i o n s i n w e i g h t o f b l u e grouse, (Dendragapus o b s c u r u s ) , Condor, 75:312-321. Sadleir,  R.M.S., 1 9 6 5 . The r e l a t i o n s h i p b e t w e e n a g o n i s t i c b e h a v i o u r and p o p u l a t i o n changes i n t h e deermouse, P e r o myscus m a n i c u l a t u s . J . A n i m . E c o l . , 1 4 : 3 3 1 - 3 5 2 .  Southwick,  C.H., 1 9 5 5 . R e g u l a t o r y mechanisms o f h o u s e mouse populations: s o c i a l behaviour a f f e c t i n g l i t t e r survival. E c o l o g y , 36:627-634.  25  S t i r l i n g , I.G., 1968. A g g r e s s i v e b e h a v i o u r and t h e d i s p e r s i o n o f female b l u e grouse. Can. J . Zoology, 46:405. S t i r l i n g , I.G., and J . F . B e n d e l l , 1970. The r e p r o d u c t i v e b e h a v i o u r of b l u e grouse. S y e s i s , 3:161-171. Svendsen, G.E., and K.B. A r m i t a g e , 1973. M i r r o r - i m a g e s t i m u l a t i o n a p p l i e d t o f i e l d b e h a v i o u r a l s t u d i e s . E c o l . , V o l . 54, No. 3, pp. 623-627. Theberge, J . , 1971. P o p u l a t i o n f l u c t u a t i o n and changes i n t h e q u a l i t y o f Rock P t a r m i g a n i n A l a s k a . Unpubl. Ph.D. t h e s i s , U n i v e r s i t y o f B.C. Watson, A., 1964. A g g r e s s i o n and p o p u l a t i o n r e g u l a t i o n i n r e d g r o u s e . N a t u r e , 202 (4931) :506-507. Watson, A., and J e n k i n s , D., 1968. Experiments on P o p u l a t i o n C o n t r o l by T e r r i t o r i a l B e h a v i o u r i n Red Grouse. J . Anim. E c o l . 37, 595-614. Watson, A., and R. Moss, 1969. Dominance, s p a c i n g b e h a v i o u r and aggression i n r e l a t i o n t o population l i m i t a t i o n i n v e r t e b r a t e s . I n : Animal P o p u l a t i o n s i n R e l a t i o n t o T h e i r Food Resources. The B r i t i s h E c o l o g i c a l S o c i e t y , Aberdeen, March 1969. pp. 167-222. Z w i c k e l , F.C., 1965. E a r l y m o r t a l i t y and t h e numbers o f b l u e grouse. Unpubl. PhD. t h e s i s , U n i v e r s i t y o f B.C. 153 pp. Z w i c k e l , F.C., and J . F . B e n d e l l , 1967a. A snare f o r c a p t u r i n g b l u e grouse. J . W i l d l . Mgmt., 31:202-204. Z w i c k e l , F.C., and J.F. B e n d e l l , 1967b. E a r l y m o r t a l i t y and t h e r e g u l a t i o n o f numbers i n b l u e g r o u s e . Can. J . Z o o l . , 45:815-851. Z w i c k e l , F.C., 1972. Removal and r e p o p u l a t i o n o f b l u e grouse i n an i n c r e a s i n g p o p u l a t i o n . J . W i l d l . Mgmt. 36, pp. 1141-1152.  

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