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Moose-wolf dynamics and the natural regulation of moose populations Messier, François 1984

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MOOSE-WOLF DYNAMICS AND THE NATURAL REGULATION OF MOOSE  POPULATIONS  By Francois  Messier  B.Sc,  University  Laval,  1976  M.Sc,  University  Laval,  1979  A THESIS SUBMITTED  IN PARTIAL  THE REQUIREMENTS DOCTOR OF  FULFILMENT  FOR THE DEGREE OF PHILOSOPHY in  THE FACULTY OF GRADUATE STUDIES (Department  we a c c e p t  of Animal  this  to the  Science)  t h e s i s as c o n f o r m i n g  required  standard  THE UNIVERSITY OF BRITISH COLUMBIA September © F.  1984  Messier,  1984  OF  )E-6  In  presenting  requirements  this f o r an  of  British  it  freely available  agree for  that  f o r reference  understood  that  be  h i s or her copying  f i n a n c i a l gain  fulfilment at the  I  study.  copying by  representatives.  allowed  Frangois  of  Animal  Sciences  The U n i v e r s i t y o f B r i t i s h 1956 Main M a l l Vancouver, Canada V6T 1Y3  Date  (3/81)  20 S e p t .  1984  of  of  Columbia  this  thesis  of  my  It i s  this  without  MESSIER  make  further  the head  permission.  Department  University  and  granted  n o t be  the  shall  or publication  shall  of  the Library  f o r extensive  p u r p o s e s may  o r by  degree  I agree that  permission  scholarly  in partial  advanced  Columbia,  department  for  thesis  thesis my  written  ABSTRACT  Long t e r m d a t a  indicate  populations  s o u t h w e s t e r n Quebec s t a b i l i z e a t  in  =0.40 a n i m a l - k m .  To t e s t  - 2  competition,  that  naturally  equilibrium density  (0.37=H)  (=0.23=M,  M e a s u r e m e n t s of  breadth,  heart  weight,  and a t  1982.  poorer  at  2 lower  T h e r e was no i n d i c a t i o n t h a t  competition To t e s t  for  population regulation  d y n a m i c s were s t u d i e d a t data  in area L ) .  individuals, km , 2  In  the a v a i l a b l e  between area,  combat,  were c o l l e c t e d  of  greater  use of  densities.  regulatory  alternative  in areas  was that  by p r e d a t o r s , moose-wolf  s i z e s averaged  territories  respectively.  (partial 5.7  and  3.7  390 and 255  r e p r e s e n t e d «*0% and The w o l f  population  s u f f e r e d from a h i g h e r intraspecific  feeding observations food r e s o u r c e s at  E a c h pack k i l l e d on a v e r a g e  moose*100 d a y s  1981  process.  success in producing pups.  summer s c a t s and w i n t e r  from  s e a s o n s of  body c o n d i t i o n  r a t e due t o m a l n u t r i t i o n and l e t h a l  and f r o m a l o w e r  this  cranial  a r e a s H and M, p a c k s a v e r a g e d  i n a r e a M, as compared t o a r e a H, mortality  near  same 3 moose d e n s i t i e s  year-long t e r r i t o r y  and i n t e r s t i c e s  30% of  the  forage  and hence no e v i d e n c e  forage e x p l a i n s the  of  densities  head l e n g t h ,  and k i d n e y w e i g h t  t h e h i g h moose d e n s i t y ,  by  investigated  443 moose k i l l e d d u r i n g t h e autumn h a r v e s t i n g and  a density  population regulation  moose body c o n d i t i o n was  0.17=L).  r e g u l a t e d moose  H, M, and L,  5.3,  Analysis  indicated a  lower 1.8,  respectively.  moose and  1.1  January  wolf  densities  animals*100  were r e s p e c t i v e l y  km" .  of I  are  wolf  is  the p o s t n a t a l  conclude that  offers  removal  presented.  I  have an  the  (<0.5-1.0  natural  2  density-dependent)  but  effect  populations  that are  stable  those  competition.  the  regulatory  animal'km" )  than  at  effect.  be  f r o m 6.1  to  effect  r e g u l a t e d at  wolf  results  of a  hypothesis.  predation  low moose  a depensatory  higher  for  of moose p o p u l a t i o n s  that at  Quebec  where c o m p e t i t i o n  H support t h i s  concept  regulated  southwestern  Preliminary  regulation  also at  in  a density  in area  support  important  rates proved to  moose p o p u l a t i o n s  experiment of  0.36  populations.  no d e t r i m e n t a l  A review  and  i n c r e a s i n g w i t h moose d e n s i t y  r e g u l a t e d by p r e d a t o r s  forage  0.82,  Year-long predation  2  density-dependent, 19.3%  1.38,  densities  (inversely  densities.  by p r e d a t o r s  Moose  appear  high densities  can  by  t o be more  forage  iv  RESUME  L e s d o n n e e s o b t e n u e s au c o u r s d ' u n e demontrent ouest  que l e s p o p u l a t i o n s d ' o r i g n a u x  du Quebec a t t e i g n e n t  approchent  un e q u i l i b r e  0.40 a n i m a l • k m " .  phenomene,  la nourriture  l e sud-  leurs si  d i s p o n i b l e peut  de 0 . 3 7 ( H ) ,  =*0.23 (M) , e t  L e s mesures de l o n g u e u r de t e t e , et p o i d s des r e i n s , a l a chasse  n'indiquent  densites  la  expliquer  ce  largeur  furent  sportive  0.17 a n i m a l ' k m " du c r a n e ,  recueillies  en 1981 e t  a  (L) .  2  p o i d s du  s u r 443 o r i g n a u x  1982.  Les  resultats  pas une d e g r a d a t i o n de l a c o n d i t i o n p h y s i q u e d e s  animaux a l a p l u s pour  lorsque  duree  l a c o n d i t i o n physique des orignaux a ete e t u d i e e  des d e n s i t e s  abattus  habitant  A f i n de v e r i f i e r  2  c o m p e t i t i o n pour  coeur,  e t u d e de l o n g u e  forte  la nourriture  densite  d'orignaux.  n'expliquerait  La  competition  done p a s l e p r o c e s s u s de  regulat ion. Afin  de v e r i f i e r  predateurs, aux M,  les  interactions  3 memes d e n s i t e s  annuels  255 k m , e t  represents  2  =*0 e t  par m a l n u t r i t i o n  specifiques,  et  un p l u s  etudiees  d'etudes  30% de l ' a i r e  M a accuse  H et Les  moyenne de 390  les territoires  ont  disponible.  un p l u s haut  e t combats l e t h a u x  faible  les  3.7 l o u p s en moyenne.  une s u p e r f i c i e  l e s espaces e n t r e  p o p u l a t i o n de l o u p s de l ' a i r e mortalite  Dans l e s a i r e s  5.7 e t  ont c o u v e r t  respectivement  des p o p u l a t i o n s p a r  l o u p s - o r i g n a u x ont e t e  d'orignaux.  l e s meutes o n t r e g r o u p e  territoires et  la regulation  intra-  s u c c e s a p r o d u i r e des  La  taux de  louveteaux  que l a p o p u l a t i o n de l ' a i r e  excrements d ' e t e en h i v e r  et  des o b s e r v a t i o n s  a i n d i q u e une e x p l o i t a t i o n  alternatives  de n o u r r i t u r e  H.  L'analyse  d'alimentation accrue  a des d e n s i t e s  des  en moyenne 5 . 3 ,  orignaux'100  aires  En J a n v i e r , 0.82,  et  jours les  dans l e s  densites  de l o u p s ont  0.36 a n i m a u x « l 0 0  km" ,  s'est  passant  19.3% de l a  parturition. le  a  J'en  par  l'action  l'interieur  Je avoir  d'orignaux plus par  un e f f e t  (<0.5-1.0  regulateur)  le  etant  l'abondance  '1.1  respectivement.  estimees  densite  a  Le  1.38, taux  de  d'orignaux,  a des  cette  c o n c e p t que l a  2  a des  apres  faibles  Les  la dans  densites  resultats  hypothese.  par  les  regularisees  nourriture.  chez  l'orignal  p r e d a t i o n par faibles  mais un e f f e t  plus elevees.  regularisees  de  et  e x p e r i m e n t a l de l o u p s a  animal•km" ),  que c e l l e s  d'orignaux.  populations d'orignaux  regulateur  a des d e n s i t e s  stables  L,  des mecanismes de r e g u l a t i o n supporte  1.8,  p o p u l a t i o n estimee  de l ' a i r e H s u p p o r t e n t  presentee.  d'orignaux  la  des p r e d a t e u r s .  d'un c o n t r o l e  Une r e v u e  ete  sont maintenues  regulatrice  preliminaires  l o u p peut  a c c r u avec  c o n c l u s que l e s  s u d - o u e s t du Quebec  et  loups  sources  respectivement.  2  p r e d a t i o n annuel de 6.1  H, M,  des  moindres  Chaque meute de l o u p s a t u e  des  Les  est  le  densites inverse  (non-  populations  predateurs  apparaissent  a des d e n s i t e s  elevees  TABLE OF CONTENTS Page Abstract  ii  Resume  iv  T a b l e of  Contents  vi  List  of  Tables  ix  List  of  Figures  xi  Acknowledgements General  introduction  CHAPTER  I  . ..xiii 1  - BODY CONDITION AND POPULATION  REGULATION  BY  FOOD RESOURCES IN MOOSE Introduction  2  Study  4  area  Methods  7  Results  9  Evidence  of  population regulation  9  Body c o n d i t i o n vs moose d e n s i t y  13  Discussion  CHAPTER  II  19  - SOCIAL  ORGANIZATION,  AND POPULATION  SPATIAL DISTRIBUTION,  DENSITY OF WOLVES IN  TO MOOSE DENSITY  RELATION —  Introduction  24  Study  25  area  and methods  vii  Page Results  31  Mortality Social  and r e p r o d u c t i o n  31  organization  Spatial  34  distribution  37  Discussion  47  CHAPTER III  - MOOSE-WOLF  DYNAMICS  Introduction Study  areas  55 and methods  57  Results  60  Feeding ecology Age,  sex,  Killing Wolf  60  and c o n d i t i o n of  rates,  densities  food intake  dead moose rates,  and m o o s e : w o l f  Year-long predation  rates  64  and h a n d l i n g t i m e  ..  ratios  and p o p u l a t i o n b a l a n c e s  72 ....  Discussion  CHAPTER  IV  68  72 79  - MOOSE RESPONSE TO EXPERIMENTAL WOLF REMOVAL  Introduction  85  Study  87  area  and methods  Results  90  Discussion  95  Management  implications  98  viii  Page CHAPTER V - GENERAL DISCUSSION AND CONCLUSIONS  100  Literature  108  Appendix  cited  '. . 117  ix  L I S T OF TABLES Page 1.  Estimates in areas  2.  3.  o f moose d e n s i t y H, M, and L,  and annual h u n t i n g r a t e  in  f r o m 1976 t o 1983  11  A g e - s p e c i f i c head l e n g t h ( c m ) , c r a n i a l b r e a d t h ( c m ) , h e a r t w e i g h t ( k g ) , and k i d n e y w e i g h t (kg) f o r male moose, g i v e n as mean v a l u e s ± S E ( n ) , i n r e l a t i o n t o 3 l e v e l s of moose d e n s i t y i n s o u t h w e s t e r n Quebec . . . .  14  A g e - s p e c i f i c head l e n g t h ( c m ) , c r a n i a l b r e a d t h ( c m ) , h e a r t w e i g h t ( k g ) , and k i d n e y w e i g h t (kg) f o r f e m a l e moose, g i v e n a s mean v a l u e s ± S E ( n ) , i n r e l a t i o n t o 3 l e v e l s o f moose d e n s i t y i n s o u t h w e s t e r n Quebec . . . .  15  4.  A n a l y s i s of v a r i a n c e t r e a t i n g moose d e n s i t y and age e f f e c t s on head l e n g t h , c r a n i a l b r e a d t h , h e a r t w e i g h t , and k i d n e y w e i g h t o f h u n t e r - k i l l e d moose i n s o u t h w e s t e r n Q u e b e c , d a t a o f 1981 and 1982 combined 16  5.  Head l e n g t h ( c m ) , c r a n i a l b r e a d t h ( c m ) , h e a r t w e i g h t ( k g ) , a n d k i d n e y w e i g h t (kg) f o r h u n t e r - k i l l e d moose, g i v e n a s t h e p r e d i c t e d mean v a l u e s ± S E ( n ) , i n r e l a t i o n t o 3 l e v e l s o f moose d e n s i t y i n s o u t h w e s t e r n Quebec  6.  Mortality  7.  Summary o f d a t a r e g a r d i n g w o l f territories a d e q u a t e l y d e f i n e d , and t h e r e s p e c t i v e pack s i z e a n d moose d e n s i t y  38  E a r l y w i n t e r pack s i z e , w o l f and moose d e n s i t i e s r e p o r t e d f o r d i f f e r e n t b o r e a l ecosystems in North America  50  8.  9.  age)  observations  t r a c k e d d u r i n g the study  Comparative prey  M and H  10. C o m p a r a t i v e p r e y M and H  11.  f r o m 54 w o l v e s  (>3 months of  18  use by w o l v e s d u r i n g summer i n  areas  use by w o l v e s d u r i n g w i n t e r  areas  in  A g e , s e x , and m o r t a l i t y d i s t r i b u t i o n of 62 moose consumed by w o l v e s i n a r e a s M and H  32  61 63 65  X  Page 12.  Sex and age s t r u c t u r e s of moose i n h a b i t i n g a r e a s H, M, and L, as o b t a i n e d from a e r i a l s u r v e y s and autumn h u n t e r - k i l l e d animals  67  Summary of d a t a u s e d t o e s t i m a t e moose k i l l i n g r a t e by w o l v e s , and t h e t o t a l moose c o n s u m p t i o n r a t e (December t o A p r i l ) i n a r e a s H, M, and L ••••  69  14.  I n d i c e s , r e l a t i v e i n d i c e s (RI), wolf d e n s i t y (January) in areas  73  15.  D e r i v a t i o n of t h e d e n s i t y ( N o . / k m ) (>1 y r ) moose i n a r e a s H, M, and L, ( i . e . , post-natal density)  13.  16.  17.  18.  19.  20.  and b e s t e s t i m a t e s H, M, and L 2  E s t i m a t i o n of t h e and t h e y e a r - l o n g a r e a H, M, and L  of c a l f and on 1 J u n e  of  adult 75  number of moose k i l l e d by w o l v e s p r e d a t i o n r a t e i n a 1000-km 2  S u m m a r i z a t i o n of t h e r e c r u i t m e n t and m o r t a l i t y i n a r e a s H, M, and L, g i v e n as t h e p e r c e n t a g e s the p o s t - n a t a l p o p u l a t i o n s  data of  '76  77  I n d i c e s of w o l f p o p u l a t i o n s i n t h e n o n - r e m o v a l a r e a (N-R) and t h e r e m o v a l a r e a (R) b e f o r e i n i t i a t i o n of t h e w o l f r e d u c t i o n programme ( 1 9 8 0 , 1981) and d u r i n g t h e programme ( 1 9 8 2 , 1983)  91  A v e r a g e (SE) number of moose o b s e r v e d p e r 60-km plot (see methods) and e s t i m a t e s (SE) of t h e moose d e n s i t y i n t h e n o n - r e m o v a l a r e a and i n t h e r e m o v a l a r e a  93  C a l f - c o w r a t i o s i n t h e n o n - r e m o v a l a r e a and i n t h e r e m o v a l a r e a b e f o r e i n i t i a t i o n of t h e w o l f r e d u c t i o n programme ( 1 9 8 1 , 1 9 8 2 ) , and d u r i n g t h e programme ( 1 9 8 3 , 1984)  94  2  xi  L I S T OF FIGURES Page 1.  Locations  of  2.  P e r c e n t s u c c e s s of h u n t i n g p a r t i e s (3 p e r s o n s ) t o k i l l 1 moose ( t h e bag l i m i t ) i n a r e a H from 1967 t o 1982  was c o n d u c t e d  areas  H, M, and L,  Location  4.  G r o u p i n g of  5.  December t o March c h a n g e s i n t h e p r o p o r t i o n of members f o r m i n g t h e pack c o r e ( i . e . , g r o u p c o h e s i v e n e s s ) when p a c k s were o b s e r v e d i n t h e LP or i n t h e HP a r e a  or  the in  radio-tracking  of  Depiction  7.  D e p i c t i o n of t h e t e r r i t o r y s e t t l e m e n t HP a r e a s from t h e p o o l e d d a t a between 1984  8.  .9.  10.  individual  the  areas  used a n n u a l l y  packs d u r i n g the  study  5  where most  travelled  6.  by  of  area  w o l v e s was c o n d u c t e d  w o l v e s when t h e y  t h e HP a r e a  the  study  3.  of  and d e s c r i p t i o n of  where t h e  in  ........  the  (May  to  period  LP  26 35  36  April)  i n t h e LP 1980 and  10  39  and 41  S p a c e - u s e p a t t e r n s of p a c k s H6 and L4 d u r i n g t h e summer p e r i o d (May t o N o v e m b e r ) , as compared t o t h e w i n t e r p e r i o d (December t o A p r i l )  43  S p a c e - u s e p a t t e r n s of 2 n o n - r e p r o d u c t i v e p a c k s (L5 and H8) d u r i n g t h e summer p e r i o d , and t h e r e p r e s e n t a t i o n of t h e - s e a s o n a l t e r r i t o r y shift d e m o n s t r a t e d by t h e pack H2  44  Cumulative r e l a t i v e frequencies d a i l y d i s t a n c e s of r e p r o d u c t i v e p a c k s d u r i n g t h e summer  46  f o r the i n - t e r r i t o r y and n o n - r e p r o d u c t i v e  11.  C u m u l a t i v e r e l a t i v e f r e q u e n c i e s f o r the i n - t e r r i t o r y d a i l y d i s t a n c e s of p a c k s d u r i n g t h e w i n t e r p e r i o d . . .  12.  Bone marrow  d i e d of  wolf  fat  content  predation  by month f o r  moose  or m a l n u t r i t i o n  which  48 66  xii  Page 13.  Time s p e n t a t as a f u n c t i o n (pack s i z e )  a d u l t moose k i l l s ( h a n d l i n g t i m e ) o f the number of w o l v e s p r e s e n t  14. D e s c r i p t i o n of t h e s t u d y a r e a f o r t h e w o l f r e m o v a l e x p e r i m e n t , and l o c a t i o n s of t h e w o l v e s k i l l e d between November 1981 and M a r c h 1984 15.  P e r c e n t s u r v i v a l of c a l v e s a s s o c i a t e d w i t h r a d i o c o l l a r e d f e m a l e moose i n t h e w o l f r e m o v a l and t h e non-removal area  16. The p r o p o s e d c o n c e p t u a l model of food supply i n t e r a c t i o n s  predator-moose-  71  88  96 101  xi i i  ACKNOWLEDGEMENTS  Drs.  D.M.  counsel  Shackleton  throughout  committee,  Drs.  and F . L .  the  B.D.  Bunnell  study.  O t h e r members of  Owen, R . G .  input  questions,  advice  and s u g g e s t i o n s .  field assistants,  and p a r t i c u l a r l y  C.  J.  R.  M. M a c q u a r t ,  Roy, are  Financial  Peche,  Wildlife  and l o g i s t i c  S.  Klink  F.C.A.C., D.  Seip,  (our  of  Quebec.  manuscript.  t h o s e of  support  Columbia.  Societe  Walters,  form of  L.  D.  supervisory  of  several  Giguere,  St-Hilaire,  and  for  this  du L o i s i r ,  project de l a  was  Chasse,  et  s u p p o r t came from C a n a d i a n  I  Sportsmen's was  Fund,  s u p p o r t e d by  and  Leonard  Z o o l o g i q u e de Q u e b e c , and Fonds  Shackleton,  appreciated  in the  Beaumont,  Additional  scholarships. D.M.  and C . J .  The e f f o r t s  Canadian N a t i o n a l  British  (UBC),  A.  by t h e M i n i s t e r e  Service,  University  Caron,  thesis  my  and  appreciated.  provided largely de l a  the  Peterson,  provided valuable  Caron,  to  provided guidance  F.L. V.  secretary),  Bunnell,  M. C r e t e ,  Van B a l l e n b e r g h e , gave  critical  K.L.  and P.  comments on  Parker, Mills the  1  GENERAL  It  INTRODUCTION  has been commonly c l a i m e d t h a t  with ungulate  populations  Bergerud  1980; B e r g e r u d  Peterson  and Page  regulatory predation report  et  1983).  effect rate  (e.g.,  of  al.  and 2 o t h e r s  disrupted  harvesting.  It  The n a t u r a l  study  of  (Chapter of  felt  into  the  II),  I  present  in wolf  numbers.  dynamics,  predation.  populations,  by  are  reported I  review  inspire  new l i n e s  of  density  intensive  between  first  address  III,  of  via  the  a  Subsequently social  I  in  organization determining  elaborate  regulatory  effect  on of  wolf  an o n - g o i n g w o l f  natural  w i t h summary s t a t e m e n t s  and t o  I  I).  in Chapter the  is  emerge.  importance  results  moose  food resources  (Chapter  Chapter  I  " m a n i p u l a t i o n " would a  availability,  potential In  of  the  that  Here,  interactions  d i s c u s s i n g the  c o n c l u d i n g C h a p t e r V,  thought  intricate  The p r e l i m i n a r y  experiment  density.  i n f o r m a t i o n on t h e  its  i.e.,  an e q u i l i b r i u m  by  1983;  has d e m o n s t r a t e d  population process  moose body c o n d i t i o n  moose-wolf  removal  the  only  interact  1978;  al.  e q u i l i b r i u m by  population regulation  w o l v e s b e c a u s e of  changes  that  moose, and f o r a g e  q u e s t i o n of  et  regulation  1 near  from t h i s  c o m p r e h e n s i o n of  predators,  no s t u d y  with ungulate  on s u c h a s t u d y .  As an e n t r y  1974; Smuts  p r e d a t i o n upon u n g u l a t e s ,  a n a l y s e d among 3 p o p u l a t i o n s ,  clear  Keith  strongly  1983; Gasaway  Yet,  increases  was  predators  IV.  In  the  regulation  designed to research.  of  provoke  moose  2  Chapter 1  BODY CONDITION AND POPULATION  REGULATION  BY FOOD RESOURCES IN MOOSE  INTRODUCTION  Two f a c t o r s regulation Caughley  in ungulates:  mainly  because  see Macnab  these they  history  et  al.  events  accumulation, (reviews is  1980).  are and,  i n Hanks  for  intraspecific  Klein  and N o r t o n -  et  (e.g., al.  Pimlott  1983).  have been n o t a b l y to conduct  in  1981; Huot  higher  the  i n an  quality  many  affected:  scarce, field  (but  1982).  important  growth,  Among t h e  and e x p e r i m e n t a l exist  ungulate  forage  Cervidae,  there  that  due t o v a r i a t i o n s  and S t r a n d g a a r d  life-  survival  evidence  and p o p u l a t i o n d e n s i t y  (Klein  fat  r e p r o d u c t i o n and  size differences  1968; K l e i n  individuals  As a r e s u l t ,  ultimately,  food q u a n t i t y ,  1964,  preferred,  adversely  both c i r c u m s t a n t i a l  quality,  difficult  food l i m i t a t i o n ,  p o p u l a t i o n compete Kie  1980; Gasaway  (e.g.,  1983).  When f a c i n g  1970;  food resources  and p r e d a t i o n  explanations  are  of  population  1977; S i n c l a i r  1982),  1974; B e r g e r u d  testing  explain  availability  Sinclair  1982; H o u s t o n  Keith  Studies  commonly i n v o k e d t o  1970,1976;  Griffiths 1967;  are  in  food  (Verme 1963;  1972; Verme and Ozoga  3  1980a;  Suttie  et  1983; S k o g l a n d  al.  fat  1980; L e a d e r - W i l l i a m s  a c c u m u l a t i o n s and t h e i r  g o v e r n e d by t h e (Dauphine Huot  1982).  In  growth b e f o r e mortality, poor al.  short,  time  appear  al.  plane  of  1969,  Initial  1980; S k o g l a n d  premise  in  are  has a for  1983).  the  all This  study.  assessment  a more a p p r o p r i a t e a p p r o a c h w i t h w h i c h  b e c a u s e body c o n d i t i o n r e p r e s e n t s an  alces)  a  et  the p r e s e n t  habitat,  This  p e r i o d of  retardation  habitat  of  delay  in a s i z e d i f f e r e n c e  1977; S u t t i e important  growth  evaluate  evaluation  the  1979; T h o r n e  l i s t e d above c o n c l u d e d t h a t  body c o n d i t i o n i s  the  c o n s e q u e n c e s of  Many of of  1980b;  perinatal  an  authors  largely  development,  permanent e f f e c t - i s the  that  1978; R o b b i n s and R o b b i n s 1979;  which can r e s u l t  (Peterson  are  rate,  increase  1963,  and H a r p e r 1980).  evidence  in a s h o r t e r  and t h e  Reimers  foraging conditions  fecundity  and p o s t n a t a l  (Verme  1982;  1976; Verme and Ozoga  (which r e s u l t s  and H a m i l t o n  permanent e f f e c t  similar  t o be t h e most c r i t i c a l  1976; P e t e r s o n  age c l a s s e s  is  seasonal v a r i a t i o n s  lower  the w i n t e r ) ,  nutritional  Blaxter  et  the  in p r e n a t a l  the c a l v i n g  There  summer and t h e w i n t e r  1976; M i t c h e l l  retardation in  1983).  and R i c k e t t s  c o n d i t i o n than  habitat  the  direct  evaluation 'in  of  vivo'  quality.  study attempted to t e s t r e g u l a t e d by a v a i l a b l e  whether  or  not moose  food r e s o u r c e s .  (Alces  In  s o u t h w e s t e r n Q u e b e c , n o n - h a r v e s t e d moose p o p u l a t i o n s a p p e a r stabilize al.  at  a density  1981; d a t a  to  herein).  of  ^0.40 animal-km" Elsewhere  2  (Crete  et  i n N o r t h A m e r i c a , where  to  4  p r e d a t i o n does not o c c u r , densities  of  2 to  moose p o p u l a t i o n s can  4 animals-km"  production  in e a r l y  population  ( B e r g e r u d and Manuel  a p p r o a c h was  to  winter  (0.22  15 t o  (0.37  for  2  intensity  harvest  quality  (Crete et  p r e d i c t e d that body g r o w t h  rather  al.  the  than  density  higher.  our p r e d i c t i o n p r o v e s by  food resources  expected  2 lower  were  The 3  manipulated  by a r a d i c a l  change  to  area  would  body c o n d i t i o n , areas  s h o u l d be r e d u c e d and h a b i t a t  regulation  the  high density  and p o o r e r  2 lower  The  comparison.  they  competition If  total  1981).  moose i n t h e  retardation  compared t o moose i n  because  calf  1973).  and a t  and 0.17 a n i m a l ' k m " )  by d i f f e r e n t  exibit  the  1969; B l o o d  2  in density  I  25% of  animal•knr )  populations differed  in h a b i t a t  have a  s t u d y moose body c o n d i t i o n n e a r  equilibrium density densities  of  and s t i l l  2  reach  where  quality  food  s h o u l d be  be c o r r e c t ,  population  would be s u g g e s t e d .  STUDY AREA  The 3 s t u d y productive A r e a H,  (Fig.  moose h a b i t a t  the  relatively  areas  game r e s e r v e  1939 u n t i l  a limited -  are  1964.  entry  in  the  (0.37  the  ( B r a s s a r d et  La V e r e n d r y e  Since  system.  located within  i n Quebec  h i g h moose d e n s i t y  h u n t i n g was not a l l o w e d in  1)  (13,615  al.  km ), 2  animal•km" ). 2  reserve  from i t s  1964 h a r v e s t  A r e a M (2390  most 1974). had a Sport  establishment  has been r e s t r i c t e d  km ) 2  had a  moderate  by  5  Fig.  1.  Locations  of  areas  H, M, and L,  w h e r e t h e s t u d y was  conducted.  6  moose d e n s i t y game r e s e r v e  (0.22  animal•km" ),  Pontiac  (1205  has been u n r e s t r i c t e d Area L  (1185  km )  study,  area  L had been  unlimited entry  is  sites  glauca),  tolerant yellow hills.  (B.  1.4  (Canis  birch  (A.  ranging  from 75  areas  and c r e e k s  b)  cover  spicatum)  and use of  range  the  spruce  (Acer  are  moose  (Picea (Populus Shadeand  t o p s and s i d e s of  the  hazelnut  shrub s t r a t a . information  Crete on  forage.  present  in  the  study  from a p p r o x i m a t e l y  k m " , d e p e n d i n g on t h e  lower  saccharum)  and beaked  the  species are  in  papyrifera).  s u g a r maple  densities 2  white  provided additional  predator  lupus)  species  t r e m b l i n g aspen  (Betula  c o m p r i s e most of  quality,  animals*100  an  3 study  ponds,  tree  mariana),  alleghaniensis),  (1982a,  Two l a r g e Wolf  (P.  and p a p e r  availability,  Lakes,  (Abies balsamea),  spruce  cornuta)  and J o r d a n  under  with elevations  The d o m i n a n t  M o u n t a i n maple  (Corylus  harvested  present  of m i x e d c o n i f e r o u s and d e c i d u o u s  hardwoods, mainly  birch  (0.17  distributed.  balsam f i r  tremuloides),  1978.  p e r i o d p r e c e d i n g the  c o m p a r a b l e among t h e  consists  black  Hunting  Snow a c c u m u l a t i o n a v e r a g e s  unpubl.).  (Rowe 1 9 7 2 ) . are  level.  and i s  common and w e l l  stands  1978.  low moose d e n s i t y  intensively  sea  1971; M e s s i e r  The f o r e s t  15-year  rolling hills  600 m above  cm i n F e b r u a r y , (Soucy  in  system.  The r e l i e f 200 t o  abolished  2  former  and h i g h t h r o u g h o u t a r e a M s i n c e  D u r i n g the  2  km )  had a r e l a t i v e l y  2  animal•km" ).  and encompasses t h e  2  moose d e n s i t y  0.4  area. to  (Messier  7  and C r e t e are  in p r e s s ) .  unknown,  areas;  area  minimum of  but  Black  tracks  H appears  bear  (Ursus americanus)  and d r o p p i n g s a r e  to  support  0.25 b e a r » k m ~ ;  common i n  the h i g h e s t  Lachapelle  2  et  densities the 3  density  al.  (a  1984).  METHODS  D a t a were c o l l e c t e d d u r i n g autumn hunter-kill September in areas  registration  to  stations.  18 O c t o b e r  M and L.  in area  and c o u n t s  Most moose d e n s i t i e s  (1979)  techniques are  visibility data  bias  were not predictive  per  kill  et  (1981).  ( C r e t e et  al.  intervals  submitted). weakness  l o n g term d a t a Changes o v e r  16 t o  from 18  24 O c t o b e r  for  the  density  figures  the e s t i m a t e  study  because  this the  in which y e a r - s p e c i f i c  a single  other  year  aerial  projects.  and  Moose  St-Hilaire  were c o r r e c t e d  submitted).  between  cementum a n n u l i .  from p a r t i a l  Results  do not c o n s i d e r the  incisor  When a e r i a l  for survey  were c a l c u l a t e d number of  and moose d e n s i t y  20-30% of I  of  of  of  moose d e n s i t i e s  relationship  c a n be as wide as  major  al.  (hunting-effort)  Confidence  and from  d e s c r i b e d by C r e t e  available,  the  al.  initiated  2  and C r e t e  compulsory  Moose were k i l l e d  were e s t i m a t e d  (minimum 500 km )  inventory  H,  1982 a t  Age was d e t e r m i n e d by a c o m b i n a t i o n of  tooth eruption patterns  surveys  1981 and  are  hunting-days  (Crete a l . not  (Crete  from  1981).  available  and  et  i m p r e c i s i o n t o be a analysis  errors  p e r i o d exceed the  relies  on  may c o m p e n s a t e . accuracy  of  these  8  data. To p r o v i d e a d d i t i o n a l 1967-82 t r e n d of success  (i.e.,  past  % of  hunting parties  The number of  moose k i l l e d  were known e x a c t l y .  comparable throughout of  (same f o r the  years)  in  dividing  the  Four  head l e n g t h  stations, (Haigh  inter-orbital  fat  et  indices,  al.  1980)  (without  fat  selected  as p r o x i m a t e  deposit;  short-term effects)  4 days  From 1979  hunter  c o u l d be  were u s e d a s  and k i d n e y  indicators the  intake  1980a).  together,  of  end of  sensitive  and Ozoga  body d e v e l o p m e n t  and a t r i a ) , the  right  to I  weight  and t h e  body c o n d i t i o n  the  plant either  growing energy  assumed t h a t  w o u l d be e f f e c t i v e  (minimum  indicators  heart  of  harvest  measures,  breadth  Two o r g a n s ,  average  at  measured at  and c r a n i a l  T h e s e two o r g a n s a p p e a r  interpreted  t o hunt  Two s k e l e t a l  c o n d i t i o n s d u r i n g the  surplus blood,  (Verme  1967 and 1978,  t h e J a n u a r y moose  were c h o s e n .  long-term e f f e c t s ) .  deposit,  remained  were computed by  easily  breadth with s k i n ) ,  environmental  (i.e.,  sum of  the  harvest.  body c o n d i t i o n  registration  the  by t h e  rates  hunting  regulations  Between  the  number of  a predetermined area.  Harvest  H,  by t h e  and t h e  r e q u i r e d and a t h i r d  the p a r t y .  p o p u l a t i o n and t h e  period.  within  harvest  Harvest  area  1 moose,  and a g u i d e were a l l o w e d  g u i d e was no l o n g e r  included  who k i l l e d  hunting parties  this  two h u n t e r s all  for  moose p o p u l a t i o n was e s t i m a t e d  bag l i m i t ) .  parties  information  the  to d e t e c t  of phase  (without  weight left)  were  (i.e., season. or  protein  4 indices, p o o r body  9  condition. D a t a were t r e a t e d ANOVA t a b l e s  were i n t e r p r e t e d  h o m o g e n e i t y of technique  by a n a l y s i s  variances  was  each  removed)  were compared by t h e  method i s  conservative,  probability for  (i.e.,  level  means of  level  was  m a l e s and 4 f o r  variance  after  by t h e of  the  Layard  0.01.  The  age e f f e c t  Scheffe  at  0.10.  Six  GENLIN).  level.  The  chi-square predicted  had been  method.  ( N e t e r and Wasserman  set  (UBC  0.05 p r o b a b i l i t y  tested  with a p r o b a b i l i t y index  at  of  Because  this  1974:479),  age c l a s s e s  the  were u s e d  females.  RESULTS  Evidence  of  From allowed  1939 u n t i l  in area  density. have  population  1964 (25 y e a r s ) ,  H and moose l i k e l y  The i n i t i a t i o n  induced a s l i g h t  since  regulation  of  decline  moose*km" .  Population  trend  hunting success  and i n  in  the  the d e n s i t y  The c r i t i c a l  point  of  population  increment  the  harvest  annual  (SD)25 m o o s e ) .  is  This  was  2  (r =0.36, 2  that  the  r a t e was  in  at  P>0.10, P>0.10,  d a t a do not  but  0.37  i n d i c a t e d by t h e  lack  of  n=l6, F i g .  2)  n=8, T a b l e  1).  show  1983 a v e r a g e d  sustainable.  not  1964 may  Assuming a d e n s i t y  1964 and  was  equilibrium  t o be s t a b l e  (r_ = 0 . 3 5 ,  whatsoever.  between  an  t h e moose p o p u l a t i o n ,  stability  estimates here  were a t  harvesting  limited harvesting  1967 t h e p o p u l a t i o n a p p e a r s 2  sport  any of  3.6%  0.37, (186 ±  10  ~  70  8  50  CO CO LU  CO  30 3  Fig.  1975  1970  2.  Percent  ( t h e bag l i m i t ) suggests  success  of  hunting parties  i n a r e a H f r o m 1967 t o  a stable  population.  1982.  1980  (3 p e r s o n s )  to k i l l  The l a c k  of trend  1 moose (p>0.10)  Table  1.  3 study  Estimates  o f moose d e n s i t y  a r e a s f r o m 1976 t o  of the harvest  in  1983.  and a n n u a l  Males  a r e a s H, M, and L,  Area H  Moose N / km  %  N / km  3  0.30  2  Area L  Hunting  Moose  %  N / km  Hunting 2  %  1976  0.41  1977  0.36  b  4  n.d.  1978  0.37  b  4  0.29c  22  0.15b  26  1979  0.39  c  3  0.26  c  25  0.22  20  1980  0.36  c  4  0.19  c  27  0.18  21  1981  0.36  4  0.25  21  0.16  23  1982  0.32  3  0.19  25  0.16  23  1983  0.37  4  0.22c  22  0.16  31  mean  0.37  3.6  dec!ining  23  0.17  24  a Crete  c  and S t - H i l a i r e ,  b Crete et  al.  n.d.  n.d. n.d.  c  unpubl.  1981  c estimated from the h u n t i n g - e f f o r t results  a  the  No d a t a = n . d .  Area M  Moose  in  r e p r e s e n t e d 66%, 47%, and 53%  respectively.  Hunting 2  hunting rate  from Tremblay,  unpubl.)  per k i l l  (Crete et  al.  1981);  12  Crete  et  al.  (1981)  t h e moose p o p u l a t i o n 0.40 a n i m a l • k m " . (1981)  correct  the  al.  equilibrium density animal-km" loss  of  10% f o r  the  the  Both e v a l u a t i o n s d u r i n g the  is  (i.e.,  2  in  a r e a M,  evaluated  2  received  (Table  so the  =18%  (i.e.,  5%).  apparently  stable  and a t  (r_ = 0 . l O ,  P>0.10,  n = 6,  harvest  rate  of  In  harvest  area  growth  3 to  L,  rate,  a maximum  in area H  lower  than  the  2  r a t e of  0.30 a n i m a l - k m "  =5% u n t i l  P<0.05, Table  1).  annual harvest harvest  of  1).  2  1983  This  rate  r a t e was  2 3 % , minus t h e  a density  of 23%  evaluated  at  population was  0.17 a n i m a l - k m "  This population  2  sustained a  1).  as e v a l u a t e d declined  increased  0.41  annually).  t h e moose p o p u l a t i o n  Table  surveys the  plus  4%  to  animal•km" ).  3 study a r e a s ,  (Caughley.1976),  as moose d e n s i t y  rate,  24% ( T a b l e  C o n s i d e r i n g the population  5%,  sustainable  decrement,  2  or  0.37,  substantially  an a n n u a l  an a v e r a g e  the h a r v e s t  0.73  in a e r i a l  t h e moose p o p u l a t i o n was a t  population  by  a maximum o f  of  et  t h e moose d e n s i t y  v s =0.40  s l o p e = - 0 . 0 l 4 5 or  1),  at  was not  (0.37  1976 and d e c l i n e d a t  (r =0.62,  by C r e t e  bias  1967-82 d e n s i t y ,  study  a density  From r e t r o - c a l c u l a t i o n ,  suggested that  present  harvesting,  at  were d i v i d e d  1964-67 h a r v e s t  equilibrium density In  given  now-known v i s i b i l i t y  submitted).  without  H would s t a b i l i z e  were 0.28 and 0.30 but  for  (Crete et  in area  that,  O r i g i n a l values  2  al.  estimated  results by the  indicate  that  sustainable  from 24% t o a maximum of  from 0.17  to  0.37  the  animal*km" . 2  4%  1 3  Clearly,  a density-dependent  responses.  Moreover,  these  p o p u l a t i o n s were c e r t a i n l y  factor results  at  equilibrium density.  that  these  3 equilibrium densities  only  that  their  population  Note  growth r a t e  I  below  their  am not a s s u m i n g  different  b e c a u s e of  (Caughley  the 3  levels  that  population  were n e c e s s a r i l y  3 p o p u l a t i o n s were a t  equilibrium density  these  imply that  different  respective  the  underlies  equal,  levels  below  the d i f f e r e n c e s  in  1976).  Body c o n d i t i o n vs moose d e n s i t y A total 1982;  of  443 moose were measured i n t h e  232 i n a r e a H,  Observations significant kidney  118 i n a r e a M, and 93 i n a r e a  in both years effect  weight.  on e i t h e r  result  of  1981 and  L.  were c o m b i n e d b e c a u s e y e a r head l e n g t h , h e a r t  The d i f f e r e n c e  b r e a d t h was t h e  autumn  between  years  in  had no  weight,  or  cranial  an u n e x p l a i n e d d i f f e r e n c e  in area  L  only. Age i n t e r a c t e d length, indicate  cranial that  significantly  breadth,  and h e a r t  m a l e s and f e m a l e s  me t o  treat  them s e p a r a t e l y .  least  until  5 1/2 y e a r s  in  females  (Tables  2 and  No i n t e r a c t i o n for  the  4 indices  demonstrates that  of  weight.  regard to  These  grow d i f f e r e n t l y  head  observations which  obliged  Body d e v e l o p m e n t c o n t i n u e d  age  i n m a l e s and u n t i l  3 1/2  at years  3).  was p r e s e n t  (Table  w i t h sex i n  4).  between moose d e n s i t y  This  the a g e - s p e c i f i c  lack  of  and age  response  increments  (i.e.,  growth  Table 2. Age-specific head length (cm), cranial breadth (cm), heart weight (kg), and kidney weight (kg) f o r male moose, given as mean values ± SE(n), i n relation to 3 levels of moose density i n southwestern Quebec: area L, 0.17/km ; area M, 0.22 / km ; area H, 0.37/km . 2  2  2  Uyr.  Calf  Nodata=n.d.  1\ y r .  3jyr.  4J y r .  > 5Jyr.  Head length Area L  40.12±1.29 (8)  57.74±0.72(26)  60.89±1.29 (8)  63.82±1.63 (5)  64.14± 1.38 (7)  64.50  Area M  43.77±1.22 (9)  58.01 ±0.94(15)  61.61 ±1.10(11)  64.83±2.11 (3)  63.63±2.11 (3)  65.25±1.82 (4)  Area H  40.50±2.58 (2)  57.99±0.57(41)  62.73±0.62(35)  64.35±0.76(23)  64.54+0.86(18)  66.87±0.58(40)  Area L  15.62± 0.47 (8)  19.11±0.26(26)  19.14+0.47 (8)  20.10±0.59 (5)  22.22±0.54 (6)  21.00  Area M  16.08±0.42(10)  19.14 + 0.33(16)  20.72 ±0.38(12)  22.50+0.76 (3)  21.40+0.76 (3)  21.87±0.66 (4)  Area H  16.25+0.94 (2)  20.78+0.21(41 )  22.06±0.22(36)  22.56+0.28(23)  22.75+0.31(18)  23.46+0.21(41)  1.49±0.07(10)  1.73±0.12 (3)  1.97  (1)  Cranial breadth (1)  Heart weight Area L  n.d.  (1)  2.12 + 0.15 (2)  Area M  0.98±0.12 (3)  1.37±0.08 (7)  1.75+0.07 (8)  2.0U0.12 (3)  2.03  Area H  0.74±0.15 (2)  1.66±0.03(37)  1.94±0.04(33)  2.10±0.05(20)  2.30±0.06(14)  0.51±0.06 (2)  0.59±0.06 (2)  (1)  n.d. 1.97 + 0.12 (3) 2.28±0.04(33)  Kidney weight • Area L  n.d.  0.46±0.03 (7)  n.d.  Area M  0.31  (1)  0.49±0.04 (5)  0.51+0.05 (4)  Area H  0.30  (1)  0.52±0.01(38)  0.61 +0.02(29)  n.d. 0.63±0.02(18)  n.d. 0.78±0.03 (9)  n.d. n.d. 0.84±0.02(28)  Table  3.  weight  Age-specific  (kg)  density  f o r female  head l e n g t h  breadth  (cm),  m o o s e , g i v e n as mean v a l u e s t S E ( n ) ,  i n s o u t h w e s t e r n Q u e b e c : a r e a L,  0 . 1 7 / k m ^ ; a r e a M,  11  Calf  Head  (cm), c r a n i a l  21  yr.  heart weight  in relation 0.22 /km?;  (kg),  to 3 levels a r e a H,  > 3|  yr.  and  kidney of  0.37/km2.  yr.  length Area L  42.31 ±1.14  (8)  55.24 ±0.81 (16)  60.93±1.87  (3)  61.94 ±0.88(13)  Area M  44.65 ±1.02(10)  57.45±0.66(24)  62.73±1.08  (9)  63.35±0.71(21)  56.87±0.62(27)  63.11 ±1.08 (9)  62.89±0.58(31)  Area H Cranial  n.d.  breadth  Area L  14.94±0.42  (8)  17.77±0.31(15)  20.33±0.69  (3)  19.17 ±0.33(13)  Area M  15.06±0.35(12)  17.52±0.27(28)  19.50 ±0.42 (8)  19.19±0.26(22)  18.77±0.23(27)  2 0 . 3 3 ±0.40 (9)  20.55±0.20(35)  Area H Heart  n.d.  weight  Area L  0.87  (1)  1.45±0.09  Area M  0.88±0.08  (6)  1.59±0.05(15)  1.68 ± 0 . 0 8 (6)  1.97±0.06(12)  Area H  n.d.  1.70±0.04(26)  2.06 ±0.07 (8)  2.05±0.04(29)  Kidney  (5)  1.86+0.14  (2)  1.83±0.09  (5)  weight  Area L  0.30  (1)  0.44±0.03  (5)  Area M  0.26  (1)  0.47±0.02  (8)  Area H  n.d.  0.53±0.02(19)  n.d.  0.67±0.05  (2)  0.51 ± 0.03 (4)  0.54±0.03  (6)  0.61±0.03  0.68±0.01(30)  (5)  moose No d a t a = n . d .  Table 4. breadth,  Analysis  o f v a r i a n c e t r e a t i n g moose d e n s i t y  heart weight,  2.1  (calf,  yr.,  £3i  U  yr.,  on h e a d l e n g t h ,  cranial  and k i d n e y w e i g h t o f h u n t e r - k i l l e d moose i n s o u t h w e s t e r n Q u e b e c ,  1981 and 1982 c o m b i n e d . Moose d e n s i t y males  and age e f f e c t s  Z\ y r . ,  3h y r . ,  = 3 levels 4J y r . ,  ( a r e a L,  \ 5£ y r . ) ,  a r e a M, and a r e a H ) .  and 4 l e v e l s  for  females  data  of  Age = 6 l e v e l s (calf,  1|  yr.,  Heart weight  Kidney  weight  df  df  yr.).  Head l e n g t h df  P*  Cranial df  breadth P  P  P  Males 2  0.296  2  0.000  2  0.000  2  0.003  5  0.000  5  0.000  5  0.000  5  0.000  Moose d . X age  j-10  0.840  10  0.103  8  0.439  4  0.359  Error  241  Moose  density  Age  245  164  132  dales Moose  density  2  0.014  2  0.000  2  0.000  2  0.000  Age  3  0.000  3  0.000  3  0.000  3  0.000  Moose d . X age  5  0.981  5  0.732  5  0.230  4  0.230  Error  * probability  160  o f F - r a t i o MS : MS e r r o r  169  104  71.  for  17  rate)  of  kidney  head l e n g t h ,  weight  cranial  were not  interpretation  here  breadth,  affected  implies  T h i s assumption  that  6 y e a r s or correct  where moose p o p u l a t i o n s a p p e a r e d t o be  stable.  effects  of moose d e n s i t y  were f o u n d i n b o t h s e x e s , length  (Table  4).  These  individual  growth  parameters  followed  populations, between in  the  rates the  were e q u a l ,  differences  but  normally  p r e d i c t e d means;  results  achieved  the  are  i.e.,  4 indices  analysed  heart  weight  H (Table  5).  breadth in  4  that,  L and H  head  although  different  over  effect).  the  differed  (The  s l o p e s of  differed.)  analogy  regression Such  c o m p a r i s o n s of the  the  age  effect  ( f r o m t h e ANOVA, T a b l e lower  value  i n m a l e s and f e m a l e s ,  females  the  indices  age c l a s s e s  by t h e  measures  in area  on t h e  the  had a s i g n i f i c a n t l y  Cranial  in area  in  elevations  food  so p r e c e d i n g  t h e means computed a f t e r  H as p r e d i c t e d . and k i d n e y  a carry  of  morphological  within  would be t h a t  has been removed from t h e None o f  indicate  same p a t t e r n  their  My  e x c e p t i o n of male  the v a r i o u s  (i.e.,  regression analysis  lines  with the  of  levels  populations  most  and  the p o s s i b l e e f f e c t  likely  Significant  is  weight,  by moose d e n s i t y .  c o m p e t i t i o n was u n i f o r m d u r i n g t h e study.  heart  were s i g n i f i c a n t l y  in  4). area  and higher  T a b l e 5.  Head l e n g t h  (cm), c r a n i a l  breadth  (cm), heart weight  t SE(n),  h u n t e r - k i l l e d moose, g i v e n as t h e p r e d i c t e d mean v a l u e s moose d e n s i t y  i n s o u t h w e s t e r n Quebec:  P r e d i c t e d mean v a l u e s  a r e t h e means o b t a i n e d  measures f o l l o w i n g the v a r i a n c e test,  a r e a L,  analysis  0.17/km ; 2  after  presented  (kg),  in relation  a r e a M,  t h e age e f f e c t in Table 4.  and k i d n e y w e i g h t to  0.22/km ; 2  (kg)  3 levels  a r e a H,  has b e e n r e m o v e d f r o m  Means w e r e c o m p a r e d by  for  of  0.37 / k m . 2  the Scheffe  « =0.10.  Area L  Area M  Area H  Homogeneous  Differing  means  means  Males Head l e n g t h  59.91 ± 0 . 5 1 ( 5 5 )  60.69±0.56(45)  60.85±0.30(159)  L,M,H  Cranial  19.84 + 0.19(54)  20.33±0.20(48)  21.66±0.11 (161)  L,M  H  Heart weight  1.78+0.05(16)  1.75+0.04(25)  1.95±0.02(139)  L,M ; L , H  none  Kidney weight  0.54±0.03(11)  0.58+0.03(10)  0.63±0.01(123)  L,M,H  none  Head l e n g t h  57.09±0.52(40)  5 9 . 0 0 ± 0 . 4 1 (64)  58.57±0.41  (67)  M,H  L  Cranial  18.25±0.19(39)  18.12±0.14(70)  19.36±0.15  (71)  L,M  H  Heart weight  1.63±0.05(13)  1.61 ± 0 . 0 3 ( 3 9 )  1.85±0.03  (63)  L,M  H  Kidney weight  0.54±0.02  0.51 ±0.02(19)  0.61 ±0.01  (54)  L,M  H  breadth  none  Females  breadth  (8)  19  DISCUSSION  The u n d e r s t a n d i n g of  population  important  ungulate  aspects  sustained y i e l d ,  of  the  at  will  r e d u c e d much below results  regulatory In  population  the  4%,  and c )  or  equilibrium density.  near  and h a r v e s t  fairly  stable,  density  area  was  not  the  H,  compared t o  the  animals 2 other  in area  d i d not d e t e c t a s s e s s e d by t h e weight,  any d i f f e r e n c e s g r o w t h of  and k i d n e y  were f o u n d w i t h i n predictions,  weight.  none of  in  general  lower  approach.  a)  lower This  lower  in the  less than  study  stressed  densities  cranial  on  the  s u r p l u s was  body d e v e l o p m e n t  but  Quebec  the  p o p u l a t i o n growth  Differences  them was  to  substantially  head l e n g t h ,  age c l a s s e s ,  present  H were n u t r i t i o n a l l y  f o o d c o m p e t i t i o n and a h i g h e r  (i.e.,  long-term data  annual  p o p u l a t i o n s at  an  population  suggested t h a t :  its  respect  zero  southwestern  than  rate b)  in  The  specifically  game r e s e r v e ,  its  (1976)  Caughley's model,  estimated non-harvested population d e n s i t y .  t e s t e d whether  less  In  mechanism i m p l i e d r a t h e r  was  to 3  conservation, Caughley  r a t e at  to C a u g h l e y ' s model,  La V e r e n d r y e  relevant  be f o o d s t r e s s e d compared t o a  the  population density  than  system.  on moose p o p u l a t i o n d y n a m i c s  contrary  is  p o p u l a t i o n dynamics w i t h  p o p u l a t i o n w i t h a growth  equilibrium)  are  ungulate  ungulate-vegetation  ungulate  management:  and p o p u l a t i o n c o n t r o l .  p r e s e n t e d a model of to  regulation  with  rate.  rate  breadth,  I  as heart  measurements  contrary  to  the  in area  H.  In  4 cases  the  20  values  were s l i g h t l y  fact  cannot  I  but  explain  significantly  readily.  e x p e r i e n c e d more f a v o u r a b l e this  benefit  prevented  There are evidence  of  first  is  that  food t r i a l that  explanations  (i.e.,  may have s e l e c t e d  b r e a d t h and k i d n e y  whereas  heart  weight  weight are  is  3 body c o n d i t i o n  p r o v i d e d a measure growth, weight  and m o r e o v e r (Haigh  structural (Table that  specifically  5).  et  al.  and t h e  this  1980).  From t h i s  condition. (unpubl.)  to detect In  consistency  of  significantly  the  affected  Quebec p o p u l a t i o n s .  that  to  to  deer  protein intake.  justification head  the  of  length  skeletal  related  3 areas,  between  to  the  of  indices,  4 indices  u s i n g the  duration  From  carcass 2  l e a d t o a common t r e n d  from t h e  study  The  white-tailed  the  well  of  demonstrated  by e n e r g y  possible degradation  a parallel  found that  is  Among t h e  2 organ-indices  the combined i n f o r m a t i o n  sufficient  but  growth  indices.  addition,  related  measure  lack  regulation).  correlated  In  the  (1980a)  affected  indices.  for  t h e wrong  Verme and O z o g a ' s f i n d i n g s were u s e d f o r these  moose  i n a r e a H,  the p o p u l a t i o n  food resource  (Oddcoileus virginianus)  intake,  that  population process  e x p e r i m e n t s , Verme and Ozoga  cranial  fawns  I  H ; a  growth.  body c o n d i t i o n as  rate approaches zero  is  foraging conditions  two a l t e r n a t i v e  poorer  in area  One e x p l a n a t i o n  was o u t w e i g h e d by a n o t h e r  population  higher  of  I  believe  was  animal  body  same i n d i c e s ,  the v e g e t a t i v e  body growth of moose from  Crete  season  different  21  Egress is  may have c o n f o u n d e d t h e  unlikely  for  the  following  Canada do not e x h i b i t 1970;  Roussel  unpubl.).  only  areas  for  calves  disperse  5 animals  large  of  al.  over  s i z e of  and  is  suspect,  this  (Gasaway e t relying  al.  eastern (Goddard  Crete  60 moose w i t h i n  From t h e s e the  possibility  migrations 1980;  (3 marked n e a r  a border)  observations,  study a r e a s , Even  which are  1980).  the 3  moved  I  if  and  assumed  that  the  would not have v i t i a t e d  and most y e a r l i n g s  been s i m i l a r  not y e t  at  results  an age  to  The c o n c l u s i o n s would have  exclusively  on t h e s e  data  (see  Table 2  3). Fertility  malnutrition time  they  Saether  of of  give  Cervidae  birth  and H a a g e n r u d  a low p r e g n a n c y  r a t e or  the  100 y e a r l i n g  cows  productivity yearling  of  1978).  to  the  level  postnatal  there calf  was no  present  of  evidence  production.  100 cows  With the a i d  the  1980; Thomas 1982;  25 y e a r s w i t h o u t  128 per  of  b r e e d i n g , and at  corpora lutea and  sport  was  35 p e r  >2 1/2 y r .  of  radio-collared  o b s e r v e d a minimum p o s t - p a r t u m  96 c a l v e s  and o l d e r  of  a r e a H,  following  (n=23),  (unpubl.)  In  low  number of  (n=132; G a u t h i e r  time  and H a m i l t o n  1983).  harvesting),  Crete  the  (Blaxter  1966 ( i . e . ,  females,  has been r e l a t e d  females at  From 1964 t o  age  directional  1975; A d d i s o n e t  This  Moose i n  3 moose p o p u l a t i o n s were d i s c r e t e .  discreteness  of  seasonal  (Crete u n p u b l . ) .  c o n s i d e r i n g the the  al.  reasons.  During radio-tracking  study a r e a s , between  et  results.  cows a t  per  100 cows >2 y r .  of  age  b r e e d i n g time c o m b i n e d ,  (i.e.,  n=26).  22  These  indices  values  reported  (Pimlott Simkin  for  other  and J o r d a n  of  area  H was  that  in area  not  increase  1970's.  in  This episodic in coniferous  moose i n a r e a  and an  the  p o p u l a t i o n g r o w t h of  increase  observation  quality.  in-yard winter  quality  the  study  areas  2,  assertion  argued  response  had c a u s e d d u r i n g the  extensive early  e n r i c h e d markedly  Table that  the  1982a).  1).  shrub  Despite  recruitment This  habitat  quality  was  l i m i t i n g moose p o p u l a t i o n s . I  conclude that,  population  growth  deterioration resources  is  of not  in  southwestern  r a t e d o e s not  Quebec,  necessarily  low moose  reflect  foraging conditions.  Availability  sufficient  natural  to  moose p o p u l a t i o n s when d e n s i t y  explain  approaches  to  disturbance.  r e s p o n d by h i g h e r  (Fig.  removed  1982b).  likely  following  in  showed  forage  (1983)  ( C r e t e and J o r d a n  H d i d not  supports the  the  fumiferama)  infestation  in d e n s i t y  is  H and  moose  They a l s o  and Peek and Eastman  stands  1973;  in areas  10% ( C r e t e and J o r d a n  habitat  to c o n i f e r s  the  p r o d u c t i o n and  moose f o r a g e  i n moose p o p u l a t i o n of  to  1973; B l o o d  investigated  budworm ( C h o r i s t o n e u r a  mortality  that,  (1976),  superior  1981).  by h a b i t a t  exceeded  al.  an a m e l i o r a t i o n  strata  and S t e v e n s  H the p r o p o r t i o n of  Peek e t  Spruce  that  or  N o r t h A m e r i c a n moose p o p u l a t i o n s  (1982a)  limited  by moose r a r e l y  an  equal  b o t h summer and w i n t e r  They m a i n t a i n e d  that  are  1974; Hauge and K e i t h  quality  not  productivity  1959; S c h l a d w e i l e r  Crete  L.  of  0.40  of  food  regulation  of  animal'knr . 2  23  This density eastern  appears  Canada  ( C r e t e et  al.  showed t h a t  (i.e.,  rate,  - 2  at  .  there  forest) et  harvesting  densities  s h o u l d not  when d e n s i t i e s  Because  I  are  c o u l d not  s t u d y does not p r o v i d e  evidence,  studies  where  the  necessary  population process to As an a l t e r n a t i v e predation  by w o l v e s  dependent  between  regulating already al.  to  and/or  0.37  population  growth  condition  these  conditions,  hypothesis,  proof  that  moose i n  is  still  this feel  to  black  is al.  is  a  rate.  b e a r s may be  suggest  that  density-  k m , and t h u s has a 2  a testable (1981)  Quebec.  not  some c o n t r o l l e d  relate  I  of  but,  certainly  need f o r is  availability  viable,  hypothesis the  the  southwestern  food r e g u l a t i o n ,  et  to  2  null  criterion  This  I  animal-km" .  0.0 and 0.4 moose p e r  capability.  the  the p o p u l a t i o n growth  p r o p o s e d by C r e t e  (1983).  the  regulate  to  in  under  0.40  falsify  continue  occurs  study).  to  moose body  that,  below  considering a l l I  in  be c e n t e r e d on i m p r o v i n g moose  hypothesis  healthy.  f r o m 0.17  that  The f o o d r e g u l a t i o n the  moose  r a t e d i m i n i s h e d from 24%  a definitive  f o o d r e s o u r c e s does not  predation  1983; t h i s  decline  would s u g g e s t  management p o l i c y  very  al.  ranging  was no i n d i c a t i o n I  where  of  r e g i m e s a l l o w e d moose p o p u l a t i o n s  Along with t h i s  deteriorated.  habitat  equilibrium density  the p o p u l a t i o n growth  stabilize  animal'km  boreal  1981; B e r g e r u d  4% when d i f f e r e n t to  t o be t h e  hypothesis,  and B e r g e r u d  et  and  24  Chapter  SOCIAL  ORGANIZATION,  II  SPATIAL DISTRIBUTION,  AND  POPULATION  DENSITY OF WOLVES IN RELATION TO MOOSE DENSITY  INTRODUCTION  The e f f e c t  of  substantial previous  wolf  predation  theoretical  r e s e a r c h has  approach:  e.g.,  and a p p l i e d  structure,  intake  rate,  wolf  and a g e n e r a l 1977; P e t e r s o n  Peterson  al.  toward  1983).  In  functional  al.  predation and K e i t h work  manipulative  food  is  been done  and experiments  lacking  wolves to  (Mech  1980;  1984; see a l s o  information  r e s p o n s e s of  basic  rate  has  underlying processes  and M e s s i e r  the  on  Macnab the  different  densities. Here,  I  present  distribution, relying wolf  solid  and n u m e r i c a l  of  of  descriptive  selectivity.,  little  by means of  1983; C r e t e  particular,  prey  1977; F u l l e r  Relatively  relationships  (Gasaway e t  moose  1984b).  on a  Much of  numbers and t h e i r  evaluation  understanding c e r t a i n  functional  wolf  food h a b i t s ,  1966; Haber et  of  a question  interest.  focussed p r i m a r i l y  estimation  social  on moose r e m a i n s  on s o c i a l  and p o p u l a t i o n d e n s i t y  primarily  numerical  data  on moose as p r e y .  r e s p o n s e and t h e  organization, of  2 wolf  The s t u d y  important  spatial  populations documents  behavioural  the  25  attributes  associated with t h i s  experienced r e l a t i v e l y exploitation, where t h e about  whereas  process.  One w o l f  h i g h moose abundance due t o the  its  naturally  low  human  second i n h a b i t e d a c o n t i g u o u s  moose p o p u l a t i o n had been r e d u c e d  half  population  regulated density  area  (manipulated) by  to  harvesting.  STUDY AREA AND METHODS  The e c o l o g y  of  southwestern  w o l v e s was  Q u e b e c , Canada  1980 t o F e b r u a r y  1984.  c o v e r e d a 3000-km Verendrye (LP)  predominant of  prey  reserve  species  Crete  1984).  harvested) Crete  moose d e n s i t y t o t h e LP  km"  2  i n LP  i n HP,  Crete  the the  et  Beaver  3)  density of  (HP)  wolf,  Moose,  was a t  - 2  density  0.4 m o o s e » k m " (1981)  (Castor  colonies«10  km  2  the  (Messier (i.e.,  i n HP  ( w i t h i n a few  - 2  i n HP,  ( C r e t e and Samson u n p u b l . ;  density  study  km)  density  and a t  in  1981).  70% and b e a v e r  15% of  from  and non-  (Messier  presented evidence  canadensis)  La  a stable  i n LP  it  La  The b o u n d a r y of  equilibrium density  al.  in  low p r e y  HP a r e a .  of  from J u n e  and d e c l i n e d d u r i n g t h e  at  represented approximately  Fig.  area  2  s e l e c t e d because  area  3).  decreased abruptly  3.5  was  t o =0.21 a n i m a l ' k m  estimated  area.  estimated at  2  of  The n a t u r a l  was  1984).  2  (Fig.  2  i n a 6400-km  77°W;  high prey  delineated  0.37 a n i m a l ' k m "  =0.25 a n i m a l " k m "  of  and a 3400-km  the  reserve  (47°N,  This area  area  2  reserve,  south of  Verendrye  investigated  and  that  from t h e was  1 colony-10 Moose the  annual  HP  26  F i g . 3. L o c a t i o n o f t h e a r e a w h e r e most o f t h e r a d i o t r a c k i n g o f w o l v e s was conducted. The number o f d e e r h a r v e s t e d p e r 100 k m i n 1982 a r e p r e s e n t e d t o i l l u s t r a t e the deer d i s t r i b u t i o n . 2  27  prey  biomass  Deer,  the  only a l t e r n a t i v e  southeast (1984) study  i n g e s t e d by w o l v e s  of  the  (Fig.  information  and C r e t e  species,  3).  were  Messier  on o t h e r  in  press).  present  and  Crete  features  of  the  area. Fifty-four  tracked  time.  wolves  locations  Each year  week  December  from to  periods.  (methodology  i n Mech  the  frequencies  15 May t o  1 April,  but  radio-  1974).  Some  were a c c u m u l a t e d d u r i n g 2000 h of of  1 November,  tracking almost  and o c c a s i o n a l l y  rarely  i n c l u d e d group s i z e ,  in other whether  i n open h a b i t a t ) ,  in  periods.  or  not  and t h e  flights  daily  all  of  3-4  the  winter  observation  animals  identity  were  remaining  75% of  Each  flying  from 15  d u r i n g the  W o l v e s were o b s e r v e d v i s u a l l y  locations,  (i.e.,  from 14 c o n t i g u o u s p a c k s were  from an a i r p l a n e  4700 w o l f  per  ungulate  study area  have p r e s e n t e d  (Messier  were  seen  marked  individuals. Mortality  rate  of  w o l v e s was c a l c u l a t e d  cumulative  radio-tracking  Successful  year-specific  a pack  increased  providing observed  that in  the  in  focal  not pair  failed  from M a r c h t o  activity  August.  reproduction)  demonstrate  focal  sites  time,  interim  the  deaths.  following  (pup r e s t i n g  or  2)  when a t  Unsuccessful  sites  from M a r c h t o  in  when:  areas) least  1)  summer, or  the  when:  following  1)  December  reproduction  was a s c e r t a i n e d  activity  r e m a i n e d as a p a i r  and t h e  the  r e p r o d u c t i o n was a s c e r t a i n e d  intervening  was c a p t u r e d a f t e r no o r  size  times  from  were  one pup (i.e.,  a pack 2)  did  when a  December.  28  I  excepted 2 instances  was  uncertain,  territory area  1 i n HP and  Wolf  the  meanings.  In  travel  the  the  stands,  Packs t h a t  in  the  number of  individuals  territory. in  number of  socially  within  pack  size  all  bias  with  winter w o l v e s were  because  s u c h as  lake  wolves  Moreover,  group s i z e  group s i z e  tended  streams,  about  40% of  represented  is  Pack  particularly  size  wolves w i t h i n  on e a c h p a c k ' s  Pack c o h e s i v e n e s s  reproductive of  in  the  pair)  at  represented a pack,  history to  The (i.e.,  the  pack c o r e  (i.e.,  the  any  time.  For  given a  restriction  15-day was  (1 May t o  30 A p r i l )  territories  period  imposed  among p e r i o d s .  The y e a r - l o n g  the  and  pack.  refers  a pack w i t h i n This  the  moved a b o u t  d i s s o c i a t e d from the  relied  observations  in  shores,  a g r o u p when t h e y  were c o n s i d e r e d as one s a m p l e . to avoid  which a l l  pack members p r e s e n t  group i n c l u d i n g the  the  were d e c i d u o u s w i t h good  interacting  prolonged observation).  index,  for  with prey.  i n c l u d e d members t e m p o r a r i l y  this  used those  Travelling  encounters  of  I  openings.  area  Travelling  important  proportion  were a s s i g n e d t o  observations)  i n open a r e a s  in winter.  of  had a  from 3 p e r s p e c t i v e s  cases,  and f o r e s t  stands  assessment  reproduction  reared.  to A p r i l )  visual  visibility  their  on  This condition occurred frequently  and r e s t  forest  all  (December  (=*70% of  clearcut  1 in LP.  pups had been  counted r e l i a b l y .  to  assertion  g r o u p i n g was a n a l y s e d  observations  winter  which the  o v e r l a p p i n g HP and LP a r e a s ,  i n which  specific  for  were  29  delineated the  by t h e m i n i m u m - c o n v e x - p o l y g o n method u s i n g 95% of  closest  locations  Territories area-curve  formed an a s y m p t o t e  satisfied  pack's  i n s t r u m e n t e d pack  40 t o  reproductive  interpreted  as e x c u r s i o n s  and s e a s o n .  (Messier  e x c l u d e d from t h e a n a l y s i s . engaged i n e x c u r s i o n s of  On a v e r a g e ,  Values  of  LP a r e a ,  pack L8)  (Messier  the  was,  to  the  areas  territory  HP a r e a ,  of  of  could  1984). of  some e x t e n t ,  the  Moose d e n s i t y a third  estimates error.  encompassing 2 areas  c o n t r i b u t e d and t h e  respective  This  100% c o n v e x - p o l y g o n s . in  relation  estimates  area  Moose d e n s i t i e s  to m i n i m i z e the y e a r - s p e c i f i c territories  arbitrary  95% c o n v e x - p o l y g o n s  s i z e were r e g r e s s e d  yearly  not  the  east  have been p r e s e n t e d and d i s c u s s e d  and C r e t e  wolves  the^95% c o n v e x - p o l y g o n  the  and f o r  than  and were  I  because  the  were  that  5 km, but  use o f  84% (±8% SD)  regression analyses  of  f i x e s more  b)  evident  condition  computational procedure).  s i z e and moose d e n s i t y .  entire  was  than  a territory  justified  corresponded to  for  of  d e t e r m i n e d by t h e  consideration  pack  less  Radio  in press  1982;  d e p e n d i n g on  territory,  excursions objectively  circumscription  method.  It  this  locations,  established  observation-  and B a r r e t t e  Normally  80 d a i l y  status  b o r d e r of  d i s c a r d these  (Messier  and Mech 1 9 8 4 ) .  after  5 km from t h e  (i.e.,  member(s).  were assumed t o be d e f i n e d when t h e  Bowen 1982; B e k o f f was  of  of  1976 a n d  were w e i g h t e d I  HP  the (used  elsewhere  Moose d e n s i t y  density.  for  were d e r i v e d  between  to  1983  estimates  by t h e  c o u l d not  from  area  evaluate  30  moose d e n s i t y analysis  within  contains  heterogeneity Wolf  of  the  inclusion  intense of  the  was b e l i e v e d  used in  the d e n s i t y  within  distance  patterns For  of  the  were i l l u s t r a t e d  this is  type the  of  simplest  multiplied  the  census  the  area  wolves to  be  it  was  sets felt  to convey  of that  a  the  locations  separated  by 20 t o 28  frequency  2 periods:  summer,  In  For  (i.e.,  distances  and w i n t e r .  presume  distances  the a c t u a l  into  signs  daily  by c u m u l a t i v e It  way  distances  d i v i d e d data  I  by t y p i c a l  behavior,  test).  I  flights,  number o f  (Kolmogorov-Smirnov reflected  in  in  the  untracked packs.  locations  to a s s e s s  2 consecutive  were t r e a t e d  During  I  LP  it  or p a c k s  noticed.  a census a r e a ,  More q u a n t i t a t i v e l y ,  between  for  the  between  undetected p a i r s be m i n i m a l .  in  estimates.  representation  information.  the  because  interstices  and s c a n n i n g d u r i n g  presence)  locations.  hours)  to  of  p a c k s were f r e q u e n t l y  partially  Space-use  visual  of  same would have been t r u e  relative  the  each year  total-area-approach  the census area  s i z e by t h e p e r c e n t a g e  (i.e.,  wolf  used the  ground p a t r o l l i n g  territories pack  I in  known r e s i d e n t  that  l i n k e d with  i n J a n u a r y was e s t i m a t e d  The p o s s i b i l i t y  census area  Thus,  moose d i s p e r s i o n .  density  territories.  directly.  an u n d e f i n e d v a r i a n c e  and t h e HP a r e a s . allows  each t e r r i t o r y  analyses  was assumed t h a t  the  travelled  1 May t o  If  daily by  wolves.  30 November,  summer, b e c a u s e w o l v e s were r a r e l y  c o n s i d e r e d e a c h pack member i n d e p e n d e n t l y .  the  2  observed, instrumented  I  31  animals  had t r a v e l l e d  together  movement was  included  distances  p a c k s were q u a n t i f i e d  of  in  the  (same l o c a t i o n s )  analyses.  In  only  winter,  exclusively  1  daily  from t h e  pack  cores. Nonparametric a s s u m p t i o n s of Sokal  if  not  probability not  parametric  and R o h l f  tailed  statistical  1981).  of  tests  All  otherwise  tests  were v i o l a t e d  tests  except  specified.  rejection  greater  were u s e d i f  (Siegel  F - t e s t s were  Tests  than  the 1956; two-  leading to a  0.05 were j u d g e d t o  be  significant.  RESULTS  Mortality  and r e p r o d u c t i o n  Wolf system.  social Thus,  represent  than  the  of  that  i n HP  (35%;  the  Table  6).  and  animals  older  exploitation  mortality  was c o m p a r a b l e  i n LP a n n u a l l y ,  but  likely  3 months) i n LP  likely  to  mortality  period.  (55%)  high  February risk,  and s h o o t i n g )  in both study areas was  are  from O c t o b e r  during this  dynamics.  r a t e was h i g h e r  e x e r t e d by man ( t r a p p i n g  exclusively  size  than  These e s t i m a t e s  took p l a c e  family  recruitment  t h e pack  when w o l v e s e x p e r i e n c e d a h i g h e r  distributed  25%  (only  of  annual m o r t a l i t y  b e c a u s e more t r a c k i n g (54%)  attributes  deaths  the  on an e x t e n d e d  in-group m o r t a l i t y  2 important  Observations indicate  organization relies  i.e., was  Human-caused (30%  over-estimated  i n HP and due t o  the  Table  6.  Mortality  Radio-tracking  observations  f r o m 54 w o l v e s  ( > 3 months o f  t i m e when w o l v e s w e r e i n s i d e t h e i r  t h e y were o u t s i d e  as  territory  is  age) t r a c k e d referred to  during the as " h o m e " ,  study. and when  "foray".  Mortality Tracking (in  time  wolf-year)  Human-caused  Natural  Area  Home  Foray  Malnutrition  Strife  Car  HP  20.2  2.2  1  0  3  LP  19.2  2.4  5  2  1  Total  39.4  4.6  6 *  2+  Trap  Gun  Home  4  0  6  3  1  7 13  12  <> t 2 pups, + 2  adults  1 yearling,  3  adults  Overal1  Foray  33  yearly  distribution  of  population  suffered  the  occurrence  higher  malnutrition  tracking  from a h i g h e r of  and l e t h a l  ground  (Table  during e x t r a - t e r r i t o r i a l malnutrition,  natural combat  W o l v e s had a h i g h e r unfamiliar  effort).  mortality  (Table  Of  Considering  the  respective  mortalities  was  not  and  LP  because  such  6).  20 m o r t a l i t i e s , (3  from t r a p p i n g ,  homogeneous  time,  (X [1]  the  2 from strife).  distribution  = 12.7, P<0.01;  2  over  7 occurred  1 from i n t r a s p e c i f i c  tracking  of  as  r a t e when t r a v e l l i n g  movements  1 car-killed,  the  rate  mortalities  mortality 6).  Thus,  of  Table  6). Direct study  counts  due t o  the  of  pups c o u l d not  secretive  nature  when most w o l v e s were o b s e r v e d not  be d i s t i n g u i s h e d  difference  in  In  of  15 p o s s i b l e  from a d u l t s  HP a r e a ,  from the  tailed  Z-test  = 2.1,  in LP,  movements but  characteristic  this  the  In  December,  the  first  reliably.  (93%).  62% o b s e r v e d  P=0.02).  typical  of  pups  in  between least  the  HP and  LP a r e a  packs.  were  a  LP  some pups  in  one-  occasions  initially  movements  14  differs  (n=21;  3 unsuccessful  resumed  could  Nonetheless,  a den a t t e n d a n c e  non-reproductive  time,  This proportion  During  wolves subsequently of  wolves.  p a c k s p r o d u c e d at  occasions  significantly  observed,  for  in  r e p r o d u c t i o n was a p p a r e n t  area. the  the  of  be c o n d u c t e d  34  Social  organization  Travelling  group s i z e  in winter  averaged  i n HP and 2.3  in LP.  G r o u p i n g of  significantly  between  t h e HP and t h e LP a r e a s  P<0.01;  from o r i g i n a l  individuals; 46% of  the  were as  Fig.  In  6 to  in  January, in  larger  the  than  the  pack  average  pack  of  pack  = 37.5,  2  of  wolf  as  the w o l v e s ;  in  3.7±0.4  Fig.  t h e HP a r e a . in  t h e HP  5.7±0.5  This estimate  P<0.01).  (n=20)  significantly in  the  end of  s i z e d e c r e a s e d by 6% i n HP and  12% i n  influence  s i z e through the  winter  to  More  n=12)  observed  From December  significantly (one-tailed  and  area.  (SE, was  4),  the  P r e y abundance d i d not of  ( 1 8 % of  s i z e averaged  average  (X [3]  w o l v e s r e p r e s e n t e d as much as  LP a r e a  t h e HP a r e a .  (t--test = 3 . 9 ,  March,  Single  individuals  differed  distribution  8 w o l v e s were o b s e r v e d  individuals  area  4).  groups encountered  frequent  g r o u p s of  frequency  wolves  2.7  the  Z-test  LP  LP.  decrement = 1.2,  P>0.10). Pack c o h e s i v e n e s s , in  the  to  0.75  higher  pack c o r e , (Fig.  5).  in press  t h e p r o p o r t i o n of  d e c l i n e d t h r o u g h the The d e c l i n e  aggressiveness  (Messier  or  b;  pack c o h e s i v e n e s s s Whitney U - t e s t [ 7 , 7 ]  within Peterson  winter  appeared to  was d e t e c t e d  al.  1984b).  between  = 25, P > 0 . 2 0 ; . F i g .  pack  breeding  to  0.90 the  season  No d i f f e r e n c e  LP and HP a r e a 5).  members  from a b o u t  be r e l a t e d  packs d u r i n g the et  the  in  (Mann-  35  50r  n =  332 LP  AREA  HP  AREA  n=251  40  Vm 30  20  10  1111111'  10  20  30  40 2-3  TRAVELLING  F i g . 4. area.  Grouping  4-5  GROUP  o f wolves when they t r a v e l l e d  Sample s i z e s  6-8 SIZE  i n t h e LP or i n the HP  r e f e r t o t h e number o f groups  observed i n w i n t e r .  F i g . 5.  December to March changes in the proportion of members forming  the pack core ( i . e . , group cohesiveness) when packs were observed in the LP or in the HP area.  37  Spatial  distribution  Thirty this  study  (i.e.,  area-curve km ,  year-long  territories  only  averaged  asymptoted; Table  390±40  substantial (SE)  km  (SE)  overlap  in LP.  2  density. size)  P>0.50;  data  other  (r=0.49,  the  in Table  h a n d , pack  t h e HP a r e a  difference per  wolf  Therefore, related  size  (£=0.14,  83 ± (SE)7  larger  pack  n=27,  km «wolf.  In  2  t o moose  to the  255±20  s i z e or moose  (territory  the  its  s i z e may have  i n pack  s i z e was c o r r e l a t e d  was a p p a r e n t l y  625  size  as compared t o  in t e r r i t o r y  and a v e r a g e d  P=0.01).  observation-  Territory  2  t o moose d e n s i t y  7)  the  in  ( e x c l u d i n g pack H1 due t o  This difference  correlated  n=27,  km .  between LP and H P ) ,  The t e r r i t o r y - a r e a  was not  which  defined  They r a n g e d from 110 t o  (SE)  i n HP  2  for  7).  320±25  km  been c a u s e d by e i t h e r  the  territories  and had a mean of  2  were a d e q u a t e l y  density  territories  difference  in  in  pack  size. E a c h HP pack a p p e a r e d t o throughout territories when I of  the  LP a r e a ,  the  fate  but  terminated  the  study  The t e r r i t o r y  moose  the  6),  pairs  turnover)  in  the  of  in January  v a n i s h e d due t o mosaic  the  In  (Fig.  these  (higher  study.  occupy  LP  as a r e s u l r t  the area of  same g e n e r a l  2 new p a i r s 4 pairs 1984.  d e a t h of  was  area  established hazardous  Probably, 1 or  some  b o t h members.  was p r o b a b l y more dynamic  low pack  s i z e and  marginal  density. Maximum e x t e n t  1981 t o  of  interstices  1984 r e p r e s e n t e d  between t e r r i t o r i e s  2 5 % , 2 0 % , 5%,  and 5% of  t h e HP  from area  38  T a b l e 7.  Summary o f d a t a r e g a r d i n g w o l f t e r r i t o r i e s  and t h e r e s p e c t i v e  Pack  I.D.  Year  pack  size  Territory  and moose d e n s i t y  (km ) 2  Pack  size  (see  adequately  defined,  methods).  Moose  (No./km ) 2  No.  fixes  HI  1980-81  305  2  0.29  53  HI  1981-82  340  6  0.27  222  HI  1982-83  575  5  0.28  245  HI  1983-84  475  6  0.27  117  H2  1980-81  425  7  0.37  163  H2  1981-82  500  6  0.34  340  H2  1982-83  625  7  0.34  409  H2  1983-84  350  3  0.37  112  H4  1981-82  215  4  0.37  87  0.37  68  H4  1982-83  210  ?  H5  1983-84  285  ?  0.37  109  H6  1982-83  430  8  0.34  166  H6  1983-84  380  7  0.34  120  0.37  37  H8  1982-83  440  ?  LI  1981-82  275  3  0.24  74  L2  1980-81  300  4  0.25  45  L2  1981-82  225  4  0.24  81  L2  1982-83  295  3  0.23  96  L2  1983-84  250  2  0.21  58  L3  1982-83  285  2  0.23  68  L4  1980-81  130  2  0.25  32  L4  1981-82  145  5  0.24  103  L4  1982-83  185  6  0.23  256  L4  1983-84  165  6  0.21  98  L5  1982-83  400  6  0.23  114  L5  1983-84  330  3  0.21  54  L6  1983-84  190  2  0.21  55  L7  1982-83  245  4  0.23  45  L8  1981-82  365  3  0.22  84  L8  1982-83  300  2  0.22  60  r  39  F i g . 6. D e p i c t i o n o f t h e a r e a s u s e d a n n u a l l y (May t o A p r i l ) by i n d i v i d u a l packs d u r i n g the study p e r i o d . E l l i p s e s represent the approximate l o c a t i o n o f p a c k s known t o be p r e s e n t b u t n o t r a d i o - t r a c k e d . The p e r c e n t a g e s a l o n g t h e b o r d e r o f a census a r e a c o r r e s p o n d t o t h e p r o p o r t i o n o f pack l o c a t i o n s w i t h i n the census a r e a .  40  ( a v e r a g e = 15%),  and 5 5 % , 5 5 % , 4 5 % , and 30% of  (average = 45%; F i g .  6).  These p e r c e n t a g e s are  o v e r e s t i m a t e d b e c a u s e not a l l adequately (Fig.  7)  each y e a r .  settlement.  infrequently these  Some t e r r i t o r y  w h i c h means t h a t ,  within  were d e f i n e d  territory  illustration  whereas  of  mosaic  the  actual  s e c t o r s were u s e d a  1-year p e r i o d ,  m o s a i c a p p e a r e d t o be s a t u r a t e d  interstices  still  area  somewhat  s e c t o r s c o u l d have been m i s s e d by c h a n c e .  territory  (Fig.  territories  The m u l t i - y e a r  provides a better  territory  t h e LP  some of  Overall,  the  i n t h e HP a r e a ,  a c c o u n t e d f o r =30% of  t h e LP  area  7). Wolf  density  1.5,  and  1.5  '84,  respectively  area  were 0 . 6 ,  Inevitably,  only  confident  there  was (see  estimated at animals-100 from t h e a r e a M)  i n J a n u a r y of  2  6).  of  only  known pack  km"  2  the In  on how t o  size,  observed January-July  of  LP  delineate were  (9%  to  be  resident  density  i n a r e a HP,  mortality  in  and  packs  values'are was  and a t  Summer d e n s i t i e s  and an assumed r e c r u i t m e n t  '83,  and t o  density  summer, w o l f  LP.  '82,  l i m i t e d number of  average  2  1.3,  (average = 0 . 8 ) .  d i d not c o n t a i n  animals-100 km" in area  2  The main c r i t e r i a  C o n s i d e r i n g the  discussion. 2.1  km"  some s u b j e c t i v i t y Fig.  1981,  1.2,  Corresponding values  0.9 wolf-100  the census a r e a  monitored each y e a r , for  0.8,  territories  that  was e s t i m a t e d a t  (average = 1.4).  1.0,  untracked packs.  retained  t h e HP a r e a  animals-100 km"  the c e n s u s a r e a s enclose  in  were  1.2 derived  i n a r e a H,  4 pups f o r  every  17% i n packs  41  I  Fig.  L  7.  Depiction  of the t e r r i t o r y  settlement  f r o m t h e p o o l e d d a t a b e t w e e n 1980 and 1 9 8 4 . territory  aerial  for  pack  w h i c h t h e p r e s e n c e was c e r t i f i e d  observations.  LP and t h e HP  The a p p r o x i m a t e  (H7) was h e l d by a n o n - r a d i o - c o l l a r e d  accessibility)  in,the  areas  ellipsoidal  (limited many t i m e s  ground by  42  which area  reproduced s u c c e s s f u l l y M).  Two w o l v e s were c o n s t a n t l y  i n p r e s s b)  and t h e y  The d e n s i t y  estimates  Space-use the  pack's  both t r a v e l l e d d i d not  patterns  of  reproductive  pups were g e n e r a l l y rendezvous s i t e . forage  radially,  packs  adults  were l e s s  travelled  freely In  without  b).  and u s e d i t  natural  it)  they for  The r e l a t i v e l y  presence  visits  either  to  (Fig.  up t o  3 weeks  low pack  movements c r e a t e d  stay  size  of  non-reproductive pups.  They  territory, the  and  pups  (Messier  their  9a,  territory  b).  markedly  When w o l v e s u s e d a  from p r e d a t i o n  tended to  to  changes  i n movements  of  pack's  8b, c ) .  p e r m i t t e d p r o l o n g e d c o n s u m p t i o n of typical  shift  packs p a t r o l l e d  (Fig.  a  some p a c k s ,  These  and young  the  summer,  den or a t  In  of moose c a r c a s s e s  resulting  mortality,  sleeping beside 1984).  noticeable  by t h e  In  with  o b v i o u s when  8a, c ) .  more h o m o g e n e o u s l y  presence  areas.  pups f o r c e d a d u l t s  d u r i n g summer.  apparent  i n f l u e n c e d pack movements moose c a r c a s s ,  of  in  (Messier  and HP  season.  was q u i t e  throughout any  62%  wolves.  the n a t a l  (Fig.  Non-reproductive  winter,  at  Yearlings  affected  extensively  occasionally in press  9c_) .  t h e LP  lone  and t h e  effect  p r o d u c e d a sudden and q u i t e (Fig.  H,  observed alone over  The s e d e n t a r i n e s s  pups were moved o c c a s i o n a l l y  wolves  in area  pack members d i f f e r e d m a r k e d l y  found e i t h e r  and t h i s  packs  include  status  monitoring reproductive  adult  (93% of  in  its  (Messier in the  or  other  vicinity and  study  Crete area  a moose c a r c a s s .  a clumped d i s t r i b u t i o n  (often  of  These winter  43  — • X  Fig.  8.  WOLF PACK TERRITORIES  *  MOOSE  WOLF LOCATIONS  •  CARRION OR GUT PILES  CARCASSES  PUP R E S T I N G A R E A S  Space-use  (May t o N o v e m b e r ) ,  patterns  o f p a c k s H6 and L4 d u r i n g t h e s u m m e r  as compared t o t h e w i n t e r  period  (December t o  period April).  44  •  WOLF PACK TERRITORIES  X  PUP R E S T I N G A R E A S  •  CARRION OR GUT PILES  A P P R O X . N O R T H E R N LIMIT O F DEER  *  MOOSE  0  DUMP  WOLF LOCATIONS  *  DEER  CARCASSES  CARCASSES  L 5 , SUMMER 1983  H8, S U M M E R  1982  II  Fig. the  9. Space-use p a t t e r n s summer p e r i o d ,  demonstrated  of 2 non-reproductive  and t h e r e p r e s e n t a t i o n  by t h e p a c k H 2 .  packs  ( L 5 and H8)  of the seasonal  territory  during shift  45  locations. Two p a c k s ,  H1 and H 2 ,  repeated a t r a d i t i o n a l  movements w h i c h can be d e s c r i b e d as a s e a s o n a l extension  (Fig.  the  of  onset  9c,  d).  winter  Typically,  and e x p l o i t e d an a r e a deer  were s i t u a t e d .  lasted  and was  r e p e a t e d a few  winter.  A similar  observed  for  offal  pack H 1 , b u t  deer  kills  exploitation  territory  their  pups w i t h i n if  any,  the  Smirnov  Likewise,  quantity  wintering  the  of  range  of  daily  distances  reproductive  t h e LP a r e a one-tailed  than  test,  (K-S  test,  (Fig.  D[623,115]  butchery area.  In  both  motivation  for  reared  inhabited  10)  by  in  very  = 0.18,  winter,  the  = 0.28,  al.  than  analysis  of  in  Summer adequate  Non-reproductive  reproductive  daily  P=0.05).  distances  P=0.04).  1984b).  P<0.01; combined data  (Kolmogorov-  = 0.10,  an  that,  more  t h e HP a r e a  season to a c q u i r e  food (Peterson et  revealed  packs t r a v e l l e d  D[421,202]  D[26,89]  t o be t h e c r i t i c a l of  the  Deer  principal  their  normally  was  P a c k s H1 and H2 a l w a y s  portion  p a c k s wandered more f r e e l y  In  1 deer  farm and  3  during  extension  n o n - r e p r o d u c t i v e p a c k s moved g r e a t e r  LP a r e a  appears  case,  times  at  deer.  in  (K-S)  to  represented  the  summer,  extensively  this  extensions.  The a n a l y s i s during  territory  least  The t r i p  were o b s e r v e d p e r i o d i c a l l y .  likely  these  the  in  were u s e d i n a d d i t i o n  packs,  few,  southeastern  territory  where a t  of  2 weeks,  of  pack H2 moved s o u t h e a s t  small concentrations 1 or  pattern  of  packs  (K-S  LP and  HP).  distances  revealed  test,  no  46  0  2  4  6  8  10  12  14  D A I L Y D I S T A N C E (km)  Fig.  10.  distances period.  Cumulative of  relative  reproductive  In t h i s  analysis,  one i n w h i c h p u p s had b e e n  frequencies  for  and n o n - r e p r o d u c t i v e the area of reared.  the  in-territory  packs d u r i n g the  affiliation  daily summer  o f e a c h p a c k was  the  47  difference 0.04,  between  P>0.20;  Fig.  travelled  similar  territory  (K-S  of  (K-S  test,  11).  inside  D[120,130]  pack members)  single  D[451,296] = pack members  or o u t s i d e  in  pack's  combined data  (i.e.,  differ  P>0.20;  the  P>0.20;  animals  d i d not  = 0.04,  test,  = 0.04,  P a c k s and s i n g l e  D[747,566]  (K-S  Dissociated,  distances  test,  LP and H P ) .  and s o l i t a r y  LP and HP p a c k s  lone  their  wolves  travelling  combined d a t a  of  LP  and  HP) .  DISCUSSION  Recently,  changes  nutritional Present levels density  status  results, of  prey  in  the  deaths  in  LP a r e a 40% l o w e r  than  in  both study  areas  in the  1983).  investigated  view.  mortality  Low moose  This  density  r a t e due  combat.  Human-caused m o r t a l i t y the  was  disparity  i.e., to  Further,  LP p a c k s had a much  explain  at  density  HP a r e a .  and i n t r a s p e c i f i c  and c a n n o t  to  population processes,  importance,  reproduction.  this  i n a wolf  from a h i g h e r  greater  1980; K e i t h  corroborate  from n a t u r a l  suffered  of  have been l i n k e d  e c o l o g y has been  resulted  from m a l n u t r i t i o n  and p r o b a b l y success  where w o l f  resulted  LP w o l v e s  density  ( P a c k a r d and Mech  abundance,  the  approximately difference  in wolf  lower  similar in  wolf  densit ies. Extra-territorial this  study,  e x c u r s i o n s were commonly w i t n e s s e d  particularly  in  the  LP a r e a  (Messier  in press  in b,  2  1.0  •  0  •  2  1  1  4  •  1  6  8  10  .  .  12  .  .  .  •  14  DAILY DISTANCE (km)  Fig.  11.  distances daily  Cumulative of  frequencies  packs d u r i n g the w i n t e r  distance  dissociated  relative  frequency  of  for  period.  s i n g l e wolves  the Also  (lone  f r o m t h e p a c k more t h a n 5 d a y s )  in-territory  daily  presented  the  animals  during the  is  o r pack  members  same p e r i o d .  49  or A p p e n d i x ) .  Food s t r e s s  extra-territorial related  deaths  of  not  (i.e.,  sufficiently  pack  social  concealed natural altered  to  invalidate  organisation.  Natural  mortality  far-reaching  dynamics.  between  size,  on pack  year-long  territories  potentially  a b r o a d e x a m i n a t i o n of pack  s i z e appears  (Table size  8).  is  wolf  t o be  rates to  abundance  the  progressively  1983).  dispersal  increases,  in the  vacant  areas,  obtain  reproductive  (lower  of  i.e., the  pack  t o moose  regulation  mosaic  the  From  within  to defend a space  density of  pack  1976,  abundance  1982; seems  s i z e may be As  prey  becomes vacancy;  given  benefit  its  and  Fig.  7)  ( P a c k a r d and Mech 1980;  Thus,  might  (Zimen  pack  pack  a non-disperser c o u l d take advantage entire  density.  pack members.  territory  1982).  a maturing wolf status  wolf  in-group recruitment  Moreover,  territory  and B a r r e t t e  interstices  in prey  which c o u l d provoke d e l a y e d d i s p e r s a l also Messier  and  understood  strategy  the  saturated  the  An i n c r e a s e  increment  (Keith  (including  r e p r o d u c t i o n had a  related  The mechanism u n d e r l y i n g  to produce a d i r e c t  description  populations across North America,  intimately  and Mech 1 9 8 0 ) .  survival  size,  influence  complex and i m p e r f e c t l y  Packard  related  territory  in  Possibly,  some p a c k s ,  the p r e c e d i n g  and f a i l u r e  effect  mortality).  g r o u p d y n a m i c s of  non-measured pup m o r t a l i t y )  Pack  hazardous  movements and may have c a u s e d some human-  human-caused m o r t a l i t y but  upon p a c k s e n t a i l e d  in which  the  rarity  of  more by t r y i n g than  to  by d i s p e r s i n g ;  of  the  it  may  ability  of  eventually  50  Table  8.  different  Early winter boreal  pack  ecosystems  moose c o n s t i t u t e d  Area  the major  Wolf/100  size,  wolf  and moose d e n s i t i e s  in North America. prey  km?  species  Pack  Only  of wolves  size  studies are  Moose/km  reported for  for  which  included.  Source  2  S.W. Quebec (LP a r e a )  0 .8  3.7  0.2  This  N.E. A l b e r t a (Athabasca River)  0 .8  4.4 #  0.3  Fuller 1980  N.W. O n t a r i o (Pukaskwa)  1 .5  4.0 #  0.4  Bergerud et a l . 1983 f r o m T a b l e 1  S.W. Quebec (HP a r e a )  1 .4  5.7  0.4  This  S. A l a s k a (Kenai P e n i n s u l a )  1 .1 - 2 . ,0  11.2  0.8  Peterson et 1984  al.  Isle Royale, (1971 - ' 8 4 )  MI  2,.0 - 9 . 5  8.6  1.0-2.2  Peterson et unpubl.  al.  Isle Royale, (1959-'70)  MI  3,.1 - 5 . .1  11.1  1.5-2.6  Peterson  # Minimum e s t i m a t e packs.  b a s e d on c u r s o r y  observations  of  study and K e i t h  study  1977  non-radio-collared  51  breed.  T h i s advantage  c o u l d be c r u c i a l  in a h i g h l y  saturated  condition. Territory size  and p r e y  significant  abundance.  t h e HP a r e a  wolf  size.  to  size  family  territory  considering Walters  et  Peterson  et  al.  I  showed t h a t  in coyotes  size.  Thus,  Bowen  latrans)  m o d e l l i n g the  al.  1981),  influence  s h o u l d not  of  size  pack  (^3)  were l a r g e r (1982)  was  in  found  positively  adjustment  s i z e t o c h a n g i n g moose d e n s i t y ,  the c o n c u r r e n t  found a  pack  territories  (Canis  by pack  (1984b)  b e c a u s e pack s i z e was g r e a t e r .  territory  related  t o be g o v e r n e d l a r g e l y  e x p o n e n t i a l r e l a t i o n s h i p between  and t e r r i t o r y  that  s i z e appears  of  without  size  be e x p e c t e d t o  (e.g.,  reflect  reality. W a l t e r s et  al.  s h o u l d be i n v e r s e l y d e p l e t i o n of use of of  within  0.2  per  moose d e n s i t i e s ungulate  territory goal:  1)  but if  as h e r e , they  they  without  to  moose d e n s i t y  the  Data here  When w o l v e s  wolf  facing  area  per  their  Wolves can a c h i e v e (many  by u s i n g e a c h c a r c a s s  low  alternative  e x p l o i t a t i o n of  i n e a c h moose a t t a c k 2)  per  are  t h e p r e s e n c e of  because  indicate  territory-area  decreases.  prevent  would recommend  seem t o m a i n t a i n a c o n s t a n t  intensify  5 days),  I  so as  size  such a r e l a t i o n ,  size.  moose a b u n d a n c e .  by p e r s e v e r i n g  day and up t o  pack  in  2  species,  individual  of  wolf  and 0.4 m o o s e « k m " ,  i n d e p e n d e n t of  territory  to prey d e n s i t y  each t e r r i t o r y .  territory-area  between  suggested that  related  the c o n f o u n d i n g e f f e c t  that is  the  prey  (1981)  this  exceeding 1  thoroughly,  3)  52  by  f e e d i n g on a l t e r n a t i v e  resources 3),  (Messier  and 4)  on a v e r a g e  last  either  that  greater  per wolf  moose d e n s i t y .  At  approximately  85 km  Peninsula, of  of  At  2  Peterson  10-90 k m - w o l f . 2  al.  et  al.  Thus,  (1981)  is  2,  territory  t h e LP a r e a  which  In  LP p a c k s were  (winter) size  to  daily  was supports  area  and  at  the  greater  range of moose d e n s i t i e s ,  study).  Peterson  55 k m « w o l f .  W a l t e r s et  or e q u a l  of  related  et  At  al.  1.0-2.6  0.8 m o o s e « k m "  (1984)  f o u n d an  moose-km  (Isle  -2  (1977 and u n p u b l . ) the  inverse  tenable  to  occupied  2  Alaska),  Michigan),  distances  a p p e a r s t o be i n v e r s e l y  (this  1,  more i n t e n s e l y .  0.2-0.4 moose-km" , each wolf 2  food  statements  were s e a r c h i n g e a c h u n i t  Across a large  territory-area  daily  However, in  for  territory  (summer)  35% s m a l l e r  wolves  intensity.  figure  in press  statement,  of HP p a c k s .  approximately fact  and C r e t e  by p a t r o l l i n g t h e i r  regard to the  distances  and p o s s i b l y s e c o n d c h o i c e  2  (Kenai  average Royale,  reported a  range  r e l a t i o n s h i p p r o p o s e d by  when t h e  effect  of  pack  size  is considered. Present  results  may r e p r e s e n t subsist  2  perish  b).  wolves  of  a density  older  than 2)  1 year  In of  0.2  moose«km~  the  reproduce p o o r l y , in press b),  t o an u n s t a b l e  LP a r e a  age t e n d t o :  excursions frequently  which l e a d  of  t h r e s h o l d below w h i c h a pack  pack commonly ( M e s s i e r  extra-territorial All  that  reproduce s u c c e s s f u l l y .  from m a l n u t r i t i o n ,  from t h e i r in  the d e n s i t y  and/or  moose•km" ),  indicate  3)  cannot (-0.23 1)  dissociate  and 4)  (Messier  social  2  in  engage press  organization.  In  53  winter, which that  LP p a c k s u s e d an a v e r a g e  is of  far  below  15 t o  c o n d u c t e d at Keith  20 moose p e r  1980; P e t e r s o n  (3.7),  the  less  stable  than  in  The h a b i t a t  moose*km"  within  of  t h e m a r g i n a l pack  were p r e s e n t  interstices  additional  territories.  population  facilitated  by l a r g e r  (less  HP w o l f  territory  being  filled  The g r e a t e r the  density  2  reported  review very  in Keith  close  animal*km"  to 2  term d a t a ) ;  1983).  again,  of  different 0.37 be  from l a r g e r  than  rather  a b u n d a n c e of  1.4  t h e HP moose territories  animals*100 k m " , a  wolf-ungulate  2  of  1 to  and C r e t e  a relatively  studies  (Table  1984; from a n a l y s i s  8;  stable  equilibrium density  low d e n s i t y  low  10 w o l v e s - 1 0 0  The HP moose p o p u l a t i o n was  the e s t i m a t e d n a t u r a l  (Messier  territories.  packs.  accounted for  i n many o t h e r  area;  mosaic appeared to  d e n s i t y " compared t o t h e p o s s i b l e d e n s i t y km"  interstices  use of more e x t e n s i v e  food-constrained)  LP  packs.  was not n o t i c e a b l y At a d e n s i t y  8).  i n the  total of  and  the  area), the  studies  relatively  certain  enlargement  surrounding areas. the  (LP  (=30% of  the  interstices  saturated;  size  presumably with  2  and  in Table  m o s a i c a p p e a r e d t o be  =0.23 moose*km~  (HP a r e a ) ,  2  (6.2)  1977; F u l l e r  1984b; moose d e n s i t y  and d i d not p r e v e n t  from t h a t  (Peterson  f o r m a t i o n and d i s a p p e a r a n c e of of  100 days  f o u n d i n t h e HP a r e a  t h e HP a r e a ,  territories  7),  al.  territory  At a d e n s i t y between  et  moose p e r  100 d a y s o b s e r v e d i n o t h e r  c o n s e q u e n c e of  area  occasional  3.2  h i g h e r moose d e n s i t i e s  As a g e n e r a l  Fig.  the average  of  of  of  and 0.4  long-  compared t o t h e 2  54  to  4 moose«kirr  2  o b s e r v e d where p r e d a t i o n  ( B e r g e r u d and Manuel with Pimlott  (1967)  wolf-ungulate at  low  prey  Bergerud  1969; B l o o d who was  systems,  if  one of  naturally  and low p r e d a t o r  1980; B e r g e r u d  et  1973). the  occur  Therefore, first  may  (also Keith  1983; Gasaway  I  concur  to propose  regulated,  densities  al.  does n o t  et  al.  that  stabilize 1974; 1983).  55  Chapter  MOOSE-WOLF  III  DYNAMICS  INTRODUCTION  Present  theory  on t h e  natural  regulation  moose p o p u l a t i o n s  is  hypothesized that  moose p o p u l a t i o n s  availability. represent they  They r e j e c t e d  a basic  (1983)  important  suggested that  mortality  factor  Both  otherwise,  would c o n s t i t u t e (1983)  (e.g.,  the  American  of  that  (1981),  (1983)  with  moose n u m b e r s ,  effect if,  extra they  mortality  agent  et  hunting)  regulatory  r e p o r t e d an example where  apparently  induced i n s t a b i l i t y  maintained  that  extrinsic  food  agent.  a wolf  population  resources Peterson  cycles.  i n a moose p o p u l a t i o n .  Crete  et  al.  and  population  s u c h a s y s t e m would be c h a r a c t e r i z e d  term p r e d a t o r - p r e y  this  a moose  caused t h i s  p r e d i c t e d that  ultimate  argued that  may p r e v e n t an  increasingly  as a moose  authors  previously,  although  productivity  and Gasaway  g r o u p s of  forage  p r e d a t i o n may  p r e d a t i o n may become an  from i n c r e a s i n g  to d e c l i n e ;  Page  al.  density-dependent  population  notion  factor  depensatory)  population decreases. inversely  fluctuate  p r e d a t i o n d i m i n i s h e s moose  W a l t e r s et  (i.e.,  North  Peek and Eastman  the  regulating  recognized that  and s u r v i v a l . al.  ambiguous.  of  (1981),  They by  long  Bergerud  et  56  al.  (1983),  predation  and M e s s i e r  and C r e t e  below  s u s t a i n e d by a v a i l a b l e  food  To r e s o l v e  these  experiments  (e.g.,  1984; M e s s i e r  Gasaway  and C r e t e  the concept  much of  present  functional  This naturally  al.  moose was  to  test  and a t  animal'km" )  created  2  the  not  (1984)  I  the  thus  and r e l a t e d  mortality  has a r e g u l a t o r y  lose  the  n o t moose were  at  Quebec,  a density Body  capability.  of  of  condition (0.37  Based  Messier  and  predation  2  Here,  by  wolves.  density-dependent  Information  was  moose  0.40 a n i m a l « k m " .  is  on  food resources of  of  0.17  harvesting.  regulation  analyse  factor  and  evidence,  approaches I  or  (0.22  availability  to e x p l a i n n a t u r a l  this  In  equilibrium density  intensive  same 3 p o p u l a t i o n s ,  show t h a t  1983).  when  1984).  densities  by more  p o p u l a t i o n s when d e n s i t y using  and M e s s i e r  southwestern  stabilize  this  2 lower  concluded that  sufficient  In  and C r e t e  near  body c o n d i t i o n d a t a  Crete  natural  w o l v e s t o c h a n g i n g moose  whether  by p r e d a t o r s .  investigated 2  is  understood.  (Messier  animal•km" )  problem  moose p o p u l a t i o n d y n a m i c s may  n o n - h a r v e s t e d moose p o p u l a t i o n s - 2  the  1983; C r e t e  r e s p o n s e s of  study attempted  =0.40 a n i m a l ' k m  or  c o m p l e x i t y and m y s t i q u e  be b e t t e r  regulated  manipulative et  of  regulate  w h i c h c o u l d be  1984; see a l s o Macnab  and n u m e r i c a l  will  density  competing hypotheses,  particular, its  the  and  resources.  a p p r o a c h e d by means of  density  proposed that  by w o l v e s may be d e n s i t y - d e p e n d e n t  moose p o p u l a t i o n s w e l l  best  (1984)  and  presented  57  includes: rates,  wolf  feeding ecology,  handling times,  prey  moose:wolf  selectivity,  ratios,  killing  and p r e d a t i o n  rates  on moose.  STUDY AREAS AND METHODS  The 3 s t u d y  areas  habitat  i n Quebec  A r e a H,  the  stable (2390 study.  of  Area L  moose d e n s i t y  of  were c o r r e c t e d of  imprecision  (13,615  47°N,  km ),  km ) 2  Moose d e n s i t y  had a r e l a t i v e l y  s h o u l d be r e a s o n a b l y  in area M during  2  stable  and C r e t e  accurate  the  1984).  because  they  ( C r e t e et  al.  submitted;  r a d i o - c o l l a r e d moose).  However,  the  year-specific  is  visibility  low and  (Messier  2  had a  2  from =0.25 t o =0.21 a n i m a l - k i r r  77°W).  bias  still  unknown.  estimates  e c o l o g y was  i n a 6400-km  2  sector  The s c a r c i t y  o b t a i n complete determined direct  1;  moose  Moose d e n s i t i e s  from 1976 t o  were d e r i v e d  1983 t o  reduce  by  the  variability.  Wolf  3).  1974; F i g .  2  for  t h e most p r o d u c t i v e  La V e r e n d r y e  0.17 a n i m a l * k m "  regressing yearly annual  al.  0.37 a n i m a l • k i r r .  (1185  Moose d e n s i t i e s  with a i d  ( B r a s s a r d et  declined  2  located within  game r e s e r v e  density km )  are  of  o v e r l a p p i n g the wolves  in area  information there.  from s c a t  observations  analysis  s t u d i e d from June  analysis in winter  in  3 study  areas  Food h a b i t s  summer  because  (May  of  us  1 and to  wolves  t o November)  to A p r i l ) . scats  1984  (Fig.  L d i d not p e r m i t  (December  was not u s e d i n w i n t e r  1980 t o F e b r u a r y  were  Scat usually  were and  58  found d u r i n g a u t o p s i e s independent  of  wolf  were c o l l e c t e d weekly  scat  Prey  scat of  forest  in  scats  1969).  was  were washed  necessary  in a  conifer  sampling  (<1  proportions  for  week and  is  Winter  in  during  occurring the  a given  i n whole  required  scat  feeding ecology  samples,  to  the  units. the  studied  biomass  et  direct time  the  attempted  r e c o r d e d the sites  presence (except  ground-investigated  of  in  obvious  within  instances  a few  days.  The  of .  beaver  (1978).  was  to count  The m a j o r i t y of  all  a wolf  70% o f - ~ o b s e r v a t i o n s ) ,  food s o u r c e s .  as  scat  al.  from  observer  (successful  such  originally  located,  present  1  data  Each  area,  of  frequency Such  No  scats  The sum of  flights.  circled  after  (4%  during telemetry  the  value  where o n l y  took p l a c e  d e s c r i b e d by F l o y d was  and a  a  t r a c e amounts.  analysis.  to evaluate  technique  in  0.10,  and g r a s s  item r e f e r s  the and  Non-food elements  twigs,  in the  of  (Adorjan  observations  feeding  old)  food item w i t h i n  nearest  71% of  dry  included  i n g e s t e d u s i n g the  animals  was  macroscopic  collection  each  The e s t i m a t i o n  needles,  expressed  presentation  to the  items  by  impressions  1-mm mesh s c r e e n .  were not  occurrence  scats  were i d e n t i f i e d  estimated  i t e m was p r e s e n t .  content)  scat  roads w h i l e t r a p p i n g  The p r o p o r t i o n of  was v i s u a l l y  stones,  Fresh  and h a i r - s c a l e  0.05 was a s s i g n e d t o  food  summer,  comparison with a reference  food items,  estimation  In  activities.  along  remains  Kolenosky  kills.  surveys.  examination, major  of  the  and (=90%)  kills)  investigor  of  were then  59  d e t e r m i n e d the  nature  characteristics, morphological (general  source or  within  food source,  of  death,  s u c h as  condition;  was d e f i n e d a s  f r e q u e n t e d the  at  the  cause  samples  physical  feeding-day  I  the  of  1 km i f  site  (from  incisors  (age)  that  wolves  involved  during 7 days.  was q u i t e  used a food source  for  1970). at  the  reliable  food  For  of  had  example,  was  the  because wolves  many c o n s e c u t i v e  a  5 wolves  Evaluation  A  animal  interpretation).  r e c o r d e d 35 f e e d i n g - d a y s when a pack of a moose c a r c a s s  and bone marrow  daily-location  was a p p a r e n t  track  prey  collected  following Neiland  1 wolf  it  and  assessed  located  number of  commonly  days and were  easily  observed. Moose k i l l i n g  r a t e was c a l c u l a t e d  r a d i o - t r a c k i n g p e r i o d s and t h e analysis,  I  considered only  ± 7 SD d a y s )  in which  by more t h a n  every  but weather  day  tracking.  Pack  those  none of  separated  interim  from t h e kills.  winter  the  pack  52 h (=2 d a y s ) .  In  tracking  the  this sessions  relocations I  (23  were  attempted  conditions occasionally  excursions outside  cumulative  to  fly  interrupted  study areas  wolf  were  excluded. Dead moose were c l a s s i f i e d positive hair  evidence  on t h e  inside  broken v e g e t a t i o n absence  of  of  attack  thighs  or  around the  struggling  (large  1)  wolf  kills  bloody a r e a ,  if  lower  rear  site),  or  as n a t u r a l  2)  with signs  rumen f r o z e n whole  there  was  blood-soaked  on t h e  (typically  sternum with f o l d e d l e g s ,  as:  legs,  and death  in  s u c h as c a r c a s s in  the  body,  on  low  60  bone marrow strong  and aged c a r c a s s  densities  number of  were a s s e s s e d  wolves per  and t h e  area  provide  additional  present  3 population  permanent 2)  on b a r e  ground  emitting  smell).  Wolf the  fat,  between  circuit  frequencies  percentages howling.  of  of  pack,  in both areas  the  territories  s i z e of  (Messier  pack  (48  km)  wolf  of  1)  weekly  randomly  observations  hunting-days  for  territories,  in press a ) .  i n f o r m a t i o n among a r e a s indices:  M and H from  H, M, and L,  scat  in Chapter  I  counts along a  selected  road segments,  by moose h u n t e r s ,  which hunters  heard  The m e t h o d o l o g i e s a s s o c i a t e d w i t h t h e s e  be p r e s e n t e d  To  and  3)  wolves  indices  will  IV.  RESULTS  Feeding  ecology  Summer scats in  food h a b i t s  collected  scats  in areas  H (summer  area  M, w o l v e s p a r t l y  by a g r e a t e r  at  1.0  average  was  compensated f o r  and r a c c o o n estimated at  colony«10  of  (Marmota  km  - 2  9).  Moose  of  the  lower  use of  species  snowshoe h a r e  (Procyon l o t o r ) . 3.5  higher  H and 26% i n a r e a M ) .  secondary prey  monax),  628  occurrence  5 6 % , and was c o n s i s t e n t l y  = 52% i n a r e a  exploitation  woodchuck  americanus), density  H and M ( T a b l e  r a n g e d from 24 t o  area  beaver,  were d e t e r m i n e d by a n a l y s i s  For  colonies*10 km  - 2  In  moose  such  as  (Lepus  example, in area  i n a r e a M ( C r e t e and Samson  in  H,  beaver and  unpubl.),  Table 9. Comparative prey use by wolves during summer i n areas M and H. Number of scats analysed are presented i n parentheses (years combined). May & Jun Species or  M H  food items  Jul & Aug M H  (33) (109)  Sep to Nov  Weighted  M H  (152) (228)  X  M H  (35) (71)  (220) (408)  Percent prey occurrence i n scats Adult moose  7  17  10  17  10  21  9  19  Calf moose Beaver  24  36  15  28  14  35  17  33  31  28  41  26  62  33  47  30  31  14  18  16  2  T  15  9  a  Woodchuck Snowshoe hare  3  1  4  6  T  0  2  2  0  T  8  5  6  8  5  5  4  3  4  2  6  3  5  3  61  43  51  46  58  b  cent biomass o r i g i n a l l y i n g e s t e d  0  Adult moose  46  66  Calf moose Beaver  20  19  50 17  23  26  37  21  28  18  9  18  8  12 1  3  5  3 1  7 T  21 5  8  Woodchuck  23 T T  0  1  2 T  3 4  4 1  2 4  3 1  1.0  1. 0  1.0  2..7 10,.4 12,.4  2.5  5,.7  1.8  Snowshoe hare Fruits Other items  0 3  T T 2  Relative number of prey ingested 1.0 Adult moose 1.0 5.1 3.4 Calf moose 9.2 3.3 . Beaver Woodchuck Snowshoe hare a  '  Fruits Other i t e m s r  e  1 4 5  3  1.0  1.0 2.2 3.1 5.0  1..1  3.5  5..0  29.7  4.9  2.0 8.5 12.0  6.2  2.9  6.3  1  • 1.0 i, 6 12..5  2.0 ' 3.1 0.5 0  3.2 3.0  By late August calves acquired a darker coat that resembled adult pelage.  Thus, total moose i n Sep to Nov was apportioned on the basis of the adult-calf ratio i n Jul & Aug. b  A l l scats totaled: 1.5% small rodent, 1.0% white-tailed deer, 0.5% raccoon,  and 1.0% diverse items. 0  Estimation technique for mammals as described by Floyd et a l . 1978. Assumed  whole prey weights (kg) were: adult moose (350), calf moose (30, 60, and 130, respectively), beaver (12.5), woodchuck (2.5), snowshoe hare (1.2), other mammals as Crete et a l . 1981a. Fruit percentages were halved after Andelt and Andelt 1984. d  Consumable prey weight was estimated at 90% of whole weight for a l l prey except  moose (75%) as i n Peterson et a l . 1984. e  Traces (< 1%).  62  yet,  beaver  a r e a M.  occurrence  Relative  species  in  density  the  and 2 7 % .  In  ratios  calf:adult  of  ratios  between  of  of  of  for  the  other  ingested  alternative  relative  prey  number of  moose were c o m p a r a b l e  ( 8 . 1 : 1 and 2 8 . 4 : 1  H and 47%  secondary  Moose r e p r e s e n t e d  secondary preyradult  areas  30% i n a r e a  biomass o r i g i n a l l y  M; c o r r e s p o n d i n g v a l u e s terms  was  estimates  were not a v a i l a b l e .  84% and 65% of  the  scats  prey  respectively in areas  H and  s p e c i e s were 10% prey  ingested,  in both a r e a s ,  moose d i f f e r e d  in areas  in  but  markedly  H and M,  respectively). In so  winter,  in the  area  moose p r e d o m i n a t e d as p r e y  area H than  H as compared t o  foods  (baits,  in area  offal,  M (Table  deliberately  i n t h e a r e a M (85% of 60% i n a r e a M; T a b l e and gut p i l e s )  10).  searched  I  suspect  for  these  actions  entailed  (farms,  l o g g i n g and t r a p p e r  not p e r m i t However,  few w o l v e s  more d e p e n d e n t areas.  that  of  f o o d even  the wolf  food h a b i t s  food items  than  frequently  though  such  habitations  Limited  area  in  in area M  t o human  this  more  Human-related  camps).  inhabiting  on s e c o n d a r y  10).  wolves  types  but  feeding-days  were u s e d more  hazardous approaches  a d e s c r i p t i o n of  the  species,  information in area  did L.  were p o s s i b l y in  the  2 other  Table 10.  Comparative  M and H.  Number o f f e e d i n g - d a y s  parentheses  (years  p r e y u s e by w o l v e s d u r i n g w i n t e r (see methods)  food  or  Area  items  Moose  Offal Gut  a  b  Area  a  6  3  2  21  0  4  piles  Included carcasses  25  1  of furbearer  11  14  3  animals;  areas.  I n c l u d e d dump, f a r m and b u t c h e r y  H  (693)  4  b  i n both  in  feeding-days  85  deer  baits  are presented  60  Beaver Trapper  M  (728)  (calves & adults)  White-tailed  areas  combined).  Percent  Species  in  offal.  t r a p p i n g was  permitted  64  Age,  sex,  and c o n d i t i o n  Carcasses winter  of  periods.  victims  of  dead moose'  3  62 moose were consumed by w o l v e s Ground-examination  wolves,  senescence,  of  16 d i e d  malnutrition,  revealed  from c h r o n i c disease,  that  during hunting,  by a c a r ,  not  cause  of  and  mortality  mortality age;  1 was p o a c h e d but  included  was  females.  observed  in  November  to February  starvation  in  Calves, 16%,  P>0.20), structure  of but of  late  the  kills  12)  killed 11).  The  Chronic  1 to  9 years  and 9 s e n i l e were  mortality  (>12  not  occurred  w i t h no p e r i o d o f  of  from  chronic  winter.  d i d not  living  moose  differ  from the  12; X [ 2 ]  individuals, calves  but or  still,  The  (X [2] 2  assumed  21,  = 22.7,  2  comprised  11).  areas  (respectively,  Table  were e i t h e r  animals,  (Table  between  significantly  animals  old  and o l d e r  wolf-killed  deviated  Wolves s e l e c t e d  (Table  from m a l n u t r i t i o n  prime-aged a d u l t s ,  kills  (i.e.,  3  1 was  3 instances.  Malnutrition  (Fig.  c o m b i n e d H and M d a t a ;  of  dead  study.  50%, and 34% of  structure  in  with locked antlers)  Calves  this  retrieved  7 prime-aged males  included 2 pairs  years)  uncertain  (i.e.,  injuries),  succumbed f r o m w o u n d s i n f l i c t e d N  38 were  mortality  natural  during  age = 0.78,  age  7 0 , and 9% of P<0.01).  the  prime-aged,  majority  (66%)  reproductive  moose. Bone marrow content) cases  in  fat  27% of  included  reserves  wolf-killed  2 calves,  were d e p l e t e d moose  (n=33;  3 prime m a l e s ,  (<20% Fig.  of 12).  and 6 s e n i l e  marrow These females.  Table  11.  Age,  sex,  and m o r t a l i t y  distribution  o f 62 moose consumed by w o l v e s  i n a r e a s M and H.  O b s e r v a t i o n s w e r e r e c o r d e d f r o m 15 December t o 30 M a r c h o f e a c h w i n t e r  Area  Calf  1-9  9  yr.  cf  ?  of the  M  Area  10+ y r .  9  cf  ? (>1 y r . ) . C a l f  1-9  9  ?  yr. cf  ?  study.  H  10  9  +  yr.  cf  ? (>1  ?  1  4 , 2 , 0  3 , 0 , 0  2  5  7 , 4 , 0  7 , 1 , 0  2  0  0 , 2 , 0  5 , 0 , 0  0  0  0 , 5 , 0  4 , 0 , 0  0  Other causes ^  0  0 , 2 , 0  0 , 0 , 0  2  0  0 , 0 , 1  0 , 0 , 0  0  Uncertain  1  0 , 0 , 0  1,0,0  0  0  0 , 0 , 0  0 , 1 , 0  0  A minimum o f 4 p r i m e m a l e s p e r i s h e d f r o m combat i n j u r i e s  i n a r e a H.  Predation Natural  a  mortality  a  Included 4 deaths r e l a t e d to hunting,  and 1 r o a d - k i l l  in area  H.  yr.  66  PREDATION 100  • = ADULT  o=CALF  x = M A L N UT R ITIO N  t  80  < LL 60  o or <  40  LU Z  20  o  CO  X  i •x  __L_  NOV  DEC  JAN  DATE OF  FEB  MAR  DEATH  F i g . 12. B o n e m a r r o w f a t c o n t e n t b y m o n t h f o r m o o s e w h i c h d i e d o f wolf predation or malnutrition.  Table  12.  Sex and age s t r u c t u r e s  January a e r i a l 1984 f o r  surveys  aerial  o f moose i n h a b i t i n g  and autumn h u n t e r - k i l l e d  surveys,  No.  cows  animals.  and f r o m 1978 t o 1983 f o r  From a e r i a l  per  hunting  surveys  No. c a l v e s  100 a d u l t s  a r e a s H, M, and L,  per  100 cows  as o b t a i n e d  from  P o o l e d d a t a w e r e f r o m 1981  to  samples.  From h u n t e r - k i l l e d  % calves  % 1-9 y e a r s  (N=100%)  (N)  a  animals  % 10  +  years  (N)  Area  H  63  37  19 ( 3 3 4 )  71  (592)  10  (81)  Area  M  70  41  22 ( 2 2 4 )  69  (163)  9  (22)  Area  L  63  65  29 ( 1 9 6 )  65 ( 1 3 2 )  6  (13)  a  Calculated  as t h e % o f a l l  t h e assumed % i n  living  hunter-killed  populations.  adults  X (100-% calves  observed  in surveys) =  68  Animals  that  died  had a bone marrow naturally  associated  fat  Three  level  of  exhibited  rates,  et  al.  100 w o l f - d a y s .  M, and L,  was  killing Wolves  in areas  moose t h a n in areas  killing  and L,  packs  moose was  1.1  (Table  1.7,  and  kg/wolf/day  calf  and 350 f o r  a  and cost  autumn  1.1,  (Table  0.5,  kills  for  (2-4  if  an a d u l t  13).  not  because  calves.  1 2 : 9 , and  for  packs  (5-8  Table  in 13).  lower  The r a t i o  of  in areas  1.3,  0.8,  H, and  M, 0.7  3 areas, represent  weight  moose; consumable  the  their  1:1  the  pack  for  frequently  T h e s e moose i n t a k e s whole  a  23 d a y s  Large  approximately  (assuming  moose  and by  individuals;  in  These  all,  from 8 t o  compensated  18:3,  100 p a c k -  and 0.4 moose p e r  were l o c a t e d  100 w o l f - d a y s  13).  time  moose p e r  moose c a r c a s s e s .  Overall,  respectively 1.6  and  their  M and L p a r t l y  moose were consumed p e r  previous  and h a n d l i n g  H and M s u c c e e d e d more  small  respectively.  the  that most,  4 days  r a t e by s c a v e n g i n g  killed:scavenged  in January  28 h o u r s on a v e r a g e ,  remained at  moose and from 2 t o  individuals)  to  sessions  recorded every  wolves  effort  respectively  am c o n f i d e n t  during tracking  that  adult  I  (2  prime,  phenomenon may r e f l e c t  1.8,  rates correspond respectively  fact  depletion  rates,  r a t e s were 5 . 3 , H,  12).  1982).  food intake  in areas  location  fat  with a high reproductive  Killing  killed  < 20% ( F i g .  This occasional  Clutton-Brock  Killing  days  consistently  dead males  1 in F e b r u a r y ) .  (see  from o l d age or m a i n o u r i s h m e n t  2.8,  = 150 kg f o r  fraction  = 0.75  a of  Table rate  13.  Summary o f d a t a u s e d t o e s t i m a t e moose k i l l i n g  (December t o A p r i l )  tracking  sessions  in wolf-days  (1 pack  (w.-d)  i n a r e a s H, M, and L.  and p a c k - d a y s  Area  w.-d  The a n a l y s i s  l o c a t i o n e v e r y 28 h o u r s on a v e r a g e ,  Tracking-effort  N  r a t e by w o l v e s ,  N  p.-d  and t h e t o t a l  moose  was b a s e d on d a t a c o l l e c t e d f r o m 14 p a c k s ) .  consumption  during  Tracking-effort  intensive is  expressed  (p.-d).  No. moose consumed  Moose k i l l i n g  Predation  Others  N/100 w.-d  rate  N/100  Total  p.-d  moose consumed  N/100 w.-d  N/100  H  Packs 2 - 4  577  193  6  1  1.04  3.11  1.21  3.63  1033  146  12  2  1.16  8.22  1.36  9.59  1610  339  18  3  1.12  5.31  1.30  6.19  Packs 2-4  1283  413  5  5  0.39  1.21  0.78  2.42  Packs 5-8  1293  242  7  4  0.54  2.89  0.85  4.55  All  2576  655  12  9  0.47  1.83  0.82  3.21  270  90  1  1  0.37  1.11  0.74  2.22  Packs  a  5-8  All Area  M  Area L Packs 2-4  Pack  size  p.-d  70  whole  weight;  11).  If  of  it  the prey  20% of t h e sample c o n s i s t i n g o f c a l v e s ,  i s accepted that  moose c o m p r i s e d 8 5 % , 6 0 % , and 60%  biomass  (as  ingested  areas  H and M; 60% f o r a r e a  total  intake  rate  food/wolf/day intake of  rates  i s best  (or  m i s s i n g small prey spent  significantly  (r=0.32, adult  moose  wolf-meals, kill.  the et  total al.  1984b)  kill kills  a moose  longer  observed,  size  failed  spent  the time  attacks  1 9 6 6 ; Haber was n o t  the time than  5 between  for  (n = 4 ) .  1981). the  (up t o 90% of 1977; P e t e r s o n  spent  than  t o p u r s u e and Of t h e 38 f r e s h 1 day  1 and 2 d a y s ,  and 1 r e c o r d of 5 d a y s .  to  available.  2 hours.  in less  at a  required  o b l i g e d us t o c o n s i d e r  29 had been made  interval),  the time  in Walters et a l .  reported that less  t o pack  for .calves  d i d not i n c l u d e (as  The number  p r o v i d e s a f i x e d number of  i n moose; Mech  a moose i s g e n e r a l l y  and 3 d a y s ,  a kill  the pack,  here  decreased  48.7 ± (SE)3.5 wolf-days  but s u c h i n f o r m a t i o n  (1977)  minimum t i m e  was n o t r e l a t e d  a s s o c i a t e d with  attempts  Total  time)  ( P<0.01, F i g . 13).  t i m e would have  time  Haber  that  so s m a l l e r  s u r r o u n d and k i l l  respective  estimate),  3 . 3 , 2 . 8 , and 2 . 6 kg of  (handling  and 12.5 ± ( S E ) 2 . 4  suggest  this  size  and a v e r a g e d  H a n d l i n g time  Including  at  2 for  u n d e r e s t i m a t e d due t o t h e p o s s i b i l i t y  a t moose k i l l s  (n = 2 5 ) ,  These r e s u l t s  L is a conservative  evaluated  per c a r c a s s  P>0.10)  in Table  items.  w i t h pack  feeding-days  indicated  0 . 1 1 , 0 . 0 9 , 0 . 0 8 kg/kg w o l f / d a y ) .  are l i k e l y  The t i m e  of  Table  (the  3 between 2  Seven o f t h e 9 p r o l o n g e d  71  • AREA H • AREA M 25,  20.4 - 1.75X  CO >LU  20  N  = 25  r  =-0.76  P  <  0.01  15  10  P A C K SIZE  Fig. of  13.  Time s p e n t  at  t h e number o f w o l v e s  a d u l t moose k i l l s present  (pack  (handling time)  size).  as a  function  72  attacks  were c o n d u c t e d by s m a l l  results  clearly  indicate  that  p a c k s of  2 to  5 wolves.  w o l v e s were p e r s e v e r i n g  These  i n many  attacks.  Wolf  densities Wolf  area  and m o o s e : w o l f  density  H and 0.82  in January averaged  in area M (Messier  From 3 p o p u l a t i o n e x t r a p o l a t e d at ratios  density 1.16  indices,  but  equal  in  higher  in area  1.38  km"  the  animals*100  in press a,  density 2  areas  in area  was e s t i m a t e d a t  in area M (Messier  indices  wolf  0.36 a n i m a l - 1 0 0  were a l m o s t  respectively)  ratios  L  Mooserwolf  H and M (27:1  2.07 a n i m a l s * 1 0 0  II).  was  14).  (47:1).  in press a).  L  and  28:1  In  summer,  wolf  km"  in area  H,  2  The r e l a t i v e  L s u g g e s t a summer d e n s i t y  of  in  2  or C h a p t e r  in area  (Table  km"  0.51  and  population wolf*100  km" . 2  Year-long predation To e s t i m a t e the  number of  and t h e  total  biological estimates  adult  year-long moose  the  predation yr)  postnatal  populations,  (stillborn,  malformation,  mortality  is  rate,  by w o l v e s o v e r reference  as e v a l u a t e d  balances  and c a l v e s  The b a s i c  recruitment  Perinatal  (^i  number k i l l e d  year. are  the  r a t e s and p o p u l a t i o n  after  the  believed  to  point  calculated  alive  the  on  1 June,  ensuing  for  i.e.,  perinatal  nutritional  I  all  mortality  adults  plus  mortality  deprivation,  be r e s t r i c t e d  etc.).  largely  to  the  Table 14.  Indices,  relative  indices  Weekly s c a t c o u n t s were conducted collected  in July  i n S e p t e m b e r and O c t o b e r  No.  scats  per  and b e s t e s t i m a t e s and A u g u s t  seen  per  100 h u n t i n g - d a y s RI  of w o l f  density  1980-'81-'82-'83,  (January)  i n a r e a s H, M, and  and d a t a u s e d i n t h e 2 o t h e r  L.  indices  X (No. h . - d )  RI  No. days w i t h p e r 100  Wolf  howling  hunting-days  X (No. h.-d)  density  RI  X o f RI  N o . / 100 k m  Area H  5.6  (1440)  1.00  5.1  (1172)  1.00  11.2  (1145)  1.00  1.00  1.38  Area M  4.6  (1440)  0.82  2.5  ( 483 )  0.49  4.9  ( 510 )  0.44  0.58  0.82  Area L  1.4  (1392)  0.25  1.1  ( 631 )  0.22  3.6  ( 631 )  0.32  0.26  0.36  a  Average d e n s i t y  packs,  i n January 1 9 8 0 - ' 8 1 - ' 8 2 - ' 8 3  and t e r r i t o r y  size  were  1981-'82.  No. w o l v e s  100 km • week  X ( N o . km.week)  (RI),  (Messier  submitted  e s t a b l i s h e d f r o m an i n t e n s i v e  telemetry  study of  pack  size,  a).  ^ E s t i m a t e d f r o m t h e a v e r a g e RI i n a r e a L and t h e o b s e r v e d w o l f d e n s i t y  i n a r e a H ( 0 . 2 6 X 1.38  = 0.36).  2  a  a  b  number  74  the  few d a y s  following  consequently,  its  parturition  importance  The c o m p u t a t i o n of assumptions are assumptions, somewhat  I  predation  am c e r t a i n  differed mostly  increasing  for  to  predation.  respectively. and  I  from 6.1  15 and  the  to  same  thus v a l i d a t i n g  1 9 . 3 % as moose - 2  .  Summer  represented that  predation  density predation, the  and 2% o f  in areas  the  density  75 t o 85% o f  10, 5,  adults  H, M, and  among c a l v e s  implication  being  of  a mortality  juxtaposed to In  natality-mortality  for  on  balances  the  Table the  1 June a c c o u n t e d f o r  15; C r e t e  unpubl.).  and r o a d k i l l s  confidence.  The  mortality  comments.  The  were 3 8 ,  is  limiting  17, I  revealed factors  present  on  the  3 moose p o p u l a t i o n s . 100 c a l v e s  per  100  among a r e a s  importance  of  wolf  females  ( n o t e d of  predation,  c o u l d be a s s e s s e d w i t h some  importance are  factor  other  a g e d ^2 y e a r s of' age and d i d not d i f f e r  hunting,  L,  -  r e p r o d u c t i o n and s u r v i v a l .  Recruitment  the  chosen  14%.  after  chronic  Regarding  r a t e s p r o v e d t o be  The c o r r e s p o n d i n g v a l u e s  The f u l l  Table  quantify.  associated  16.  that  3 areas,  estimated  from wolf  to  a u t h o r s may have  0.37 a n i m a l . k m  I  and,  difficult  stress  the  o r i e n t e d toward c a l v e s ,  perished annually  only  but  1974)  r a t e s and t h e  other  Year-long predation  from 0.17  year-long  22,  that  values,  a s s u m p t i o n s were a p p l i e d  dependent,  often  summarized i n T a b l e s  different  comparison.  is  (Verme  not  of  black  bear  known r e l i a b l y  predation and m e r i t  and specific  75  Table  15.  moose i n  Derivation  a r e a s H, M, and L,  (A) O v e r a l l (B)  of the d e n s i t y  density  Proportion until  of c a l f  (i.e.,  post-natal  2  on 1 J u n e  on 15 J a n  o f moose  (No./km )  and a d u l t  ( >1 y r )  density).  Area H  Area M  Area L  0 . .37  0 . ,23  0 . ,17  0..91  0 . ,94  0.,97  0 . ,337  0 . ,216  0 . ,165  0 . ,172  0 . ,118  0 . ,074  1.,00  1.,00  1.,00  0 . ,172  0 . ,118  0 . .074  0 . .51  0 . 33  0 . ,24  a  surviving  1 Jun*  (C) D e n s i t y o f  adults  on 1 J u n ,  (A) X (B) (D) D e n s i t y o f f e m a l e s (E) No. c a l v e s / f e m a l e perinatal (F)  Density of  (H) O v e r a l l  b  c  after  mortality^  density,  (D) X ( E )  (C) + ( F )  and C r e t e  These f i g u r e s  densities  >2 y r  calves,  From M e s s i e r  a g e d >2 y r  in  press.  were d e r i v e d f r o m t h e w o l f  killing  ( T a b l e 1 4 ) , and a b a s e l i n e m o r t a l i t y  ° Calculated  as  (C) X ( l -% o f c a l v e s  data from Table 12. % of y e a r l i n g s ^ A composite  on 1 J u n ,  i.e.,  on 1 J u n  among a r e a s  U  o f 4% p e r y r  the % of calves  a proportional  (see  [2] = 0.87,  text).  among  i n Jan e q u a l l e d  adults); the  mortality.  f e m a l e s were a s s o c i a t e d w i t h  (5-10 days a f t e r 2  (Table 13), wolf  i n J a n ) X (% o f f e m a l e s  s a m p l e o f 45 r a d i o - t r a c k e d  minimum o f 46 c a l v e s d i d not d i f f e r  I inferred that  rates  parturition);  P>0.50).  calf  a  production  76  Table 16.  E s t i m a t i o n of the number of moose k i l l e d by wolves and the y e a r - l o n g  p r e d a t i o n r a t e i n a lOOO-km? area H, M, and L.  The c a l c u l a t i o n was made independently  i n the summer (S=May t o Nov) and w i n t e r p e r i o d (W = Dec t o A p r ) .  Area H  Area M  S  (A) Wolf numbers  a  (B) Food i n t a k e  (kg/wolf/day)  (C) Moose consumed (kg)  W  Area L  S  W  S  W  20.7  13.8  11.6  8..2  5.1  3.,6  2.6  ...  2.2  --  2.1  -  -  500  b  0  Calf  3200  —  1180  Adult  6700  —  2510  -••-  1050  -  (D) Number moose k i l l e d ^ Calf  57.0  8.2  23.4  2..0  9.8  0..7  Adult  17.9  15.2  6.7  3..8  2.8  1..3  CE) Year-long p r e d a t i o n r a t e (%)  a  19..3  e  '  10..7  6.1  See t e x t .  * C a l c u l a t e d as the observed w i n t e r food i n t a k e (3.3, 2.8, and 2.6 kg/wolf/day  in  areas H, M, and L, r e s p e c t i v e l y ) f o r a d u l t s , and h a l f these amounts f o r pups (=40%  of  w o l v e s ) ; " o n l y the weighted food i n t a k e s are presented. c  C a l c u l a t e d as (A) X (B) X (the p r o p o r t i o n of a d u l t or c a l f moose i n the biomass  o r i g i n a l l y i n g e s t e d ; Table 9 ) .  P r o f i l e of wolf d i e t i n area L was assumed t o be s i m i l a r  t o t h a t i n area M. d In summer, we assume t h a t 10% of the c a l f biomass et a l . 1980, B a l l a r d et a l . 1981)  ( a f t e r c o n s i d e r a t i o n of Franzmann  and 30% of the a d u l t biomass represented  see a l s o notes c and d of Table 9 f o r the c a l c u l a t i o n .  In w i n t e r , the e s t i m a t i o n s were  d e r i v e d d i r e c t l y from the observed k i l l i n g r a t e per 100 w o l f - d a y s t h a t c a l v e s count f o r 35% of e  kills.  C a l c u l a t e d as (D)-j-[(H) i n Table |5 X 1000].  scavenging;  (Table 11), assuming  77  Table 17. Summarization of the r e c r u i t m e n t and m o r t a l i t y data i n areas H, M, and L, given as t h e percentages o f the p o s t - n a t a l p o p u l a t i o n s .  Values i n parentheses are  not known r e l i a b l y (see t e x t ) .  Area H  Area M  33.8  35.3  31.0  3.1  22.2  23.7  19.3  10.7  6.1  1.0  0.0  0.0  Recruitment, ( F ) ^ (H) i n Table IS  Area L  Mortality Hunting  a  Wolf p r e d a t i o n , (E) i n Table Ik Road k i l l s *  (5.0)  (2.0)  (1.5)  (4.0)  (4.0)  (4.0)  (+1.3)  (-3.6)  (-4.3)  Black bear p r e d a t i o n ° Other n a t u r a l m o r t a l i t i e s  d  Net balance Population trend, 1976-1983 Comments  e  Stable  D e c l i n i n g at ~5%per y r  Stable  P o s s i b l e egress  Possible ingress from area H  Possible from area Recruitment l i k e l y underestimated by 2-3% /  a From 1980 t o 1984, annual hunting averaged 15.9, 73.3, and 59.2 moose/1000 km i n 2  areas H, M, and L, r e s p e c t i v e l y ; h u n t e r - k i l l e d r e g i s t r a t i o n was compulsory and poaching appeared n e g l i g i b l e . b  Each y e a r , 50-70 moose (4.4/1000 km ) were k i l l e d by accident along the 117 Highway 2  t r a v e r s i n g area H. c  Assuming t h a t 15%, 10%, and 5% o f the c a l v e s were k i l l e d by black bears i n areas  H, M, and L, r e s p e c t i v e l y . d  Assuming a b a s e l i n e m o r t a l i t y of 4% f o r a l l areas (see t e x t ) .  e  From the a n a l y s i s presented by Messier and Crete ( i n p r e s s ) .  f We suspect an o v e r - r e p r e s e n t a t i o n of males i n area L (Table 12); assuming t h a t males comprised 30% of a d u l t s as i n area M ( s i m i l a r r a t e of h a r v e s t i n g ) , the r e c r u i t m e n t r a t e i s 34.3%.  78  Black  bear  predation  is  inherently  b e c a u s e moose c o n s u m p t i o n p e r small.  The p o s s i b l e  from t h e  of  high population density  375 b l a c k  May,  June,  bear  scats  and J u l y ,  (Lachapelle  of  et  from t h e  age and t h e  to p r e d a t i o n Larsen  al.  1984).  of  1984).  premise  losses  et  with  additional  20-25% of  predation  difference August  in bear  1980 t o  km,  in a l l  respectively).  that black  15%, 5%, bears  in  likely  density.  in areas areas;  the  L i m i t e d data  the  in t r a c e  et  12)  due t o scat  9.6,  efforts  In  from r a d i o - c o l l a r e d  areas  a H.  an M and  I  and  bear  1440, and  (Table  1 June  in area  in J u l y  and. 2.4  were k i l l e d  areas  on  apparent  considerations,  the c a l v e s  1981;  1 January,  respectively  = 2880,  =10 d a y s  indicates  counts  3.3,  H, M, and L,  3 respective  al.  until  the  amounts  attributable  females  which  (the  estimated  between  recruitment.  lower  in  moose  would be e x p e c t e d  (Table  From t h e s e  and 5% of  calf  (=80%)  predation  Weekly  1983 r e s u l t e d  s c a t s * 1 0 0 week-km methodology  was  of  100 c a l v e s : 1 0 0  37:100  Analysis  H d u r i n g months of  mortality mostly  carnivore  the w o l f .  1980; B a l l a r d  females  originates  facultative  remains  is  from w o l f  was  bear  this  quantify  presumably  predation  in area  calf  al.  The o b s e r v e d r a t i o  L,  is  occurred only  winter  60-65 c a l v e s : 1 0 0  l o s s of  to  Bear p r e d a t i o n may be that  Beginning  bear  s u c h as  that  killed)  following  and s u b s t r a c t i n g ratio  carnivore  revealed  (Franzmann  bear  of  collected  o n l y members p r e s u m a b l y  crudely  individual  i m p o r t a n c e of  compared t o a o b l i g a t o r y  difficult  (same 1440 week-  evaluated  e a c h summer by  17).  f e m a l e moose s u g g e s t a  79  chronic  mortalily  moose-yr).  I  chronic higher  predation  mortality, all  f o u n d at  I  values  Keith  1980; Hauge and K e i t h  i n a r e a M, and s t a b l e natality-mortality  strengthen  the  many p o s s i b l e  human  occurrence  This  estimated  figure  of  1981; Gasaway  H,  estimates,  slightly  in area  L  et  al.  17).  declining  (Messier  r e a l i s m of s o u r c e s of  the  These  with  (Rolley  and  1983).  moose p o p u l a t i o n s (=5% p e r  and C r e t e  b a l a n c e s matched r e a s o n a b l y (Table  to  chronic  conforms  c o m p a r a b l e moose d e n s i t i e s  in area  trends  relatively  or  the  tentatively  1976-83 d e n s i t y  population  ( a r e a H)  curtailed  areas.  other  appeared s t a b l e  was  d u r i n g 84  C o n s i d e r i n g t h a t m a l e s may be s u b j e c t e d  4% f o r  From t h e  2 cases  such m o r t a l i t y  ( a r e a s M and L)  natural at  (Crete unpubl.;  that  either  mortality.  mortality  2-3%  suspect  uncommon b e c a u s e exploitation  of  year)  1984).  well with  The these  correspondences  underlying estimates,  imprecision  although  remain.  DISCUSSION  In  southwestern  Quebec,  the p l a s t i c i t y  of  prey  selection  wolves  in  r e s p o n s e t o moose d e n s i t y  r e p r e s e n t e d an  aspect  of  t h e moose-wolf  Both the a n a l y s i s  summer s c a t s greater  and w i n t e r  use o f  resources  dynamics.  feeding observations  alternative,  w i t h a lower  documented i n o t h e r  probably  less  moose d e n s i t y .  wolf  studies  (Voigt  et  important of  indicated a  profitable,  Prey  by  switching al.  1976;  food has  been  Peterson  80  1977;  Shelton Killing  and P e t e r s o n rate  reflects  engaged i n c o n s u m i n g p r e y time  searching  reported that independent number of  for the  of  prey time  pack  wolves  (Holling spent  is  at  1959).  critical  t o be d a n g e r o u s l y  deceptive  for  s e a r c h i n g between addressed. kill)  pack  (n=12,  calculated  2 kills  the  s i z e of  5.7,  the average  respectively;  Therefore,  protracted  relation  study,  3.7,  time  from T a b l e  13),  for  determinant prey.  p e r i o d s without  of  the  fact  that  locating  wolves  these  (sensu For  time  H,  (Fig.  13).  between  75 d a y s .  killing  rates  through  was  the  these dead  resources.  f o o d - s o u r c e s was  were s e a r c h i n g each u n i t  I  5 5 , and 90  r  food  M,  in press a),  (19,  at  an  in areas  15 d a y s  42,  not  by s c a v e n g i n g on a l r e a d y  moose and by e x p l o i t i n g a l t e r n a t i v e Apparently,  time  amount oi time  Wolves s u r v i v e d kills  the  s i z e was  wolves  kills  8,  the  such a  Messier  the  1)  size.  13, and  between  not  predation  amount of  w i t h pack  11,  (1981)  T h i s n o t i o n may  h a n d l i n g time  and 3.0  al.  nearly  is  t o pack  respectively; of  it  warrant  in the  was a p p r o x i m a t e l y  a major  searching  in  correlated  2 0 , and 2,  consuming 2 k i l l s  time  present  a h a n d l i n g time  Considering days  the v a r i a t i o n  was  2 reasons:  p r e s e n t e d d i d not  was n e g a t i v e l y  average and L  In  of  in d e t e r m i n i n g  prove  and 2)  and t h e amount  They a r g u e d t h a t  packs.  conclusion,  is  a moose k i l l  number of  data  a predator  W a l t e r s et  the  the  time  time)  rather,  of  is  amount of  rate,  paucity  it  the  (handling  size.  that  1983).  e n h a n c e d by  area  at  the  greater  81  intensity  where  Killing (area  L)  the  (Mech  Peterson able  rates  to  et kill  numerous  (Messier  r a n g e d from 5.3 These  rates  15-20 moose*100 p a c k - d a y s  1966; P e t e r s o n  al.  1984b).  at  how c o m p l e t e leg  in winter  moose-100 p a c k - d a y s .  compared t o studies  moose were l e s s  will,  every  It  even  kill  in area  was u t i l i z e d  bones s k i n n e d down t o  the  in areas  determined  for  M and L,  wolves  was  are  that  to  quite  1980;  wolves  was a l s o  were  (disarticulated  skeleton,  As a c o n s e q u e n c e ,  kg/kg w o l f / d a y ) , low compared t o  and  the  range  studies  0 . 1 9 kg/kg w o l f / d a y ;  Peterson  1977 and u n p u b l . ;  Fuller  Keith  et  that  wolves  rate  given  relatively  in  higher  low  of  body weight adult  refractory  involved  in  Wolf w i t h the  the markedly  aid  of  is  paralleled  to the  lower  in areas  small  change  in  (i.e.,  this  H and M were d e t e r m i n e d  Wolf  The d e n s i t y  densities  i n moose d e n s i t y .  The  packs  rates  increase  killing  females  killing  to  likely  adult  were 2 o t h e r  a s s o c i a t e d w i t h the wolf  be m i n i m a l .  is  2  s i z e of  (0.10and  animal'knr .  a kill)  from 3 i n d i c e s  possible error believed  and t h e  0.4  (=26 kg f o r  radio-collared animals.  was e x t r a p o l a t e d the  males)  it  increased their  than  wolves  period after  densities  Therefore,  s t u d y would have  a moose d e n s i t y  and 30 kg f o r longer  this  1984b).  not  remarkable  on moose i n o t h e r  al.  1.1  other  preying  1980; P e t e r s o n  a).  low  in  and K e i t h  It  hooves).  moose c o n s u m p t i o n i n a r e a H ( 0 . 0 9 especially  H.  ( a r e a H)  observed  1977; F u l l e r was a p p a r e n t  in press  factors study. reliably in area  precision. density in areas  Therefore,  L  Thus,  estimates H and M the  nearly  82  2-fold difference attributable (functional wolf  to changes response)  density.  predation  in the  I  in prey  rather  suggest  that  their  that  (see  advocacy).  In  L,  in a higher  mooserwolf  a density  area  of  selection  et  density  moose«km~ ),  L  (0.17  wolf/day of  0.13  was o b s e r v e d ,  intensively  territory  was  2  to  predation analysis,  of  I  is to  a  (in  press  other  subsist  far  of  below  suggested  density reproduce  species.  and a p o r t i o n  In  =0.08 kg meat/kg the  food  requirement  reproduce s u c c e s s f u l l y ,  found evidence  so  resulting  and/or  ungulate  but  different  a)  of  the  only  (25%)  of  as  area  1 pack  L was  that  its  H.  predation 19.3% a t  rates  increased  0.37 m o o s e ' k n r . 2  r a t e s cannot  from 6 . 1 % a t Confidence  be computed due t o  and t h e p r e s e n c e  of  the  15-30% of  Even w i t h s u c h an e r r o r ,  estimates.  s o l i d evidence  that  wolf  intervals  a s s u m p t i o n s ^ The  predation  be  in  was  0.'17  complexity  r a t e s may e a s i l y  provides  reflect  was much l o w e r ,  a s s o c i a t e d with predation the  in  perspective  D u r i n g 4 summers t h a t  pups,  in area  Estimated moose'kirr  (1977).  patrolled,  produced a l i t t e r  the  which  may  1983 f o r  a food intake  kg meat/kg w o l f / d a y  p r o p o s e d by Mech  of  a b s e n c e of 2  ratios  was  rates  difference  approximated the  2  t h r e s h o l d below w h i c h a pack c a n n o t  area  al.  Messier  0.2 m o o s e - k m "  i n the  10.7%)  and k i l l i n g  i n a management  ratio.  successfully  and  range of moose d e n s i t i e s ,  Gasaway  wolf  (19.3  moose-wolf  utility  remain q u e s t i o n a b l e  rates  than a r e l a t i v e  rate across a large  imperfectly  that  predation  of  error  the  range  the  of  study  density-  83  dependent. 1)  the  Empirical  sharp d e c l i n e  in  density,  2)  at  moose d e n s i t y ,  lower  the  justifications  p o p u l a t i o n at In  killing  tendency  and 3)  southwestern  Quebec,  plateau  at  of  1984).  Wolf  =0.4 a n i m a l ' k i r r  may i n v o l v e  depressive  females bears al.  were f o u n d t o p r e y  1980)  calves also  and p r o b a b l y  i n our  density-dependent  predation  will  the  (Messier  2  entire  the  wolf  the  ratio  of  unknown.  regulatory  wolf  surveys.  Black  (Franzmann  bear If  process  37 c a l v e s : 1 0 0  a further not  Crete  dominant  example,  in Alaska  inflicted  and  regulatory  For  low  Whether or  the  the - 2  represent  on c a l v e s  remains  reinforce  resources  animal•km .  H during January  they  situation.  to  factors.  explain  observed in area  are:  n o n - h a r v e s t e d moose p o p u l a t i o n s  mechanism but  cannot  food  d i s r u p t i o n of  below =0.2  density-dependent  predation alone  assertion  r a t e w i t h d e c r e a s i n g moose  the  p r e d a t i o n appears  other  this  t o consume a l t e r n a t i v e  moose d e n s i t i e s  a density  for  reduction  predation  it  is,  effect  et  of  of  is  bear predation  on  moose p o p u l a t i o n s . In  regards  to  indication  that  population  reaches  (Messier 1982a, other  the  wolf  factors  capable  of  density  sustainable  equilibrium density  1984; C r e t e  Overall,  limiting  limitation,  there  foraging conditions deteriorate  and C r e t e  b) .  food resource  regulating  et  al.  predation,  on r e c r u i t m e n t  of  no  when a moose  0.40  1981; C r e t e  animal'kirr and J o r d a n  in combination with and m o r t a l i t y ,  a moose p o p u l a t i o n w e l l  by a v a i l a b l e  is  below  food r e s o u r c e s .  I  the  appears the conclude  2  84  that  naturally  Quebec a r e  heavily  time wolves are on t h e  r e g u l a t e d moose p o p u l a t i o n s  in  c o n s t r a i n e d by p r e d a t o r s ,  nutritionally  abundance of m o o s e .  s t r e s s e d due t o  southwestern but a t their  the  same  own a c t i o n  85  Chapter  IV  MOOSE RESPONSE TO EXPERIMENTAL WOLF REMOVAL  INTRODUCTION  In  southwestern  ecosystems, stabilize Bergerud  Q u e b e c , as  et  a density al.  of  Habitat  investigated  near  Crete  this  natural  and J o r d a n  studied  from an a n i m a l  perspective  was not  regulation  predation,  the  resources. dependent  boreal  that  Crete  moose  condition habitat  of  quality  1982; M e s s i e r  to e x p l a i n  been  r e d u c e d by  Body  al.  lower  and  availability the  of  observed  to t e s t  limiting  whether factors  or not  c a n h o l d a moose p o p u l a t i o n  that  wolf  predation  0.0 and 0.4 m o o s e - k m  -2  is  wolf  on  can be s u s t a i n e d by a v a i l a b l e  postulated  between  1981;  of moose p o p u l a t i o n s .  combined w i t h other  I  al.  use by moose have  to a s s e s s  et  sufficient  ongoing study attempts  density  1984;  1982a, b ) .  (Crete  to  (Crete et  c o n c l u s i o n was t h a t  r e p r o d u c t i o n and s u r v i v a l , below  2  populations  i n an e f f o r t  The g e n e r a l  food resources  This  in  e q u i l i b r i u m and a t  (i.e.,  moose was a l s o  1984).  and C r e t e  p r o d u c t i o n and i t s  for comparison  harvesting;  natural  places  =0.4 a n i m a l « k n r  1983; M e s s i e r  submitted).  Crete  some o t h e r  n o n - h a r v e s t e d moose p o p u l a t i o n s a p p e a r  at  densities  in  well  food  density-  ( C r e t e et  al.  1981;  86  Messier  and C r e t e  recruitment  1984,  in p r e s s ) .  and i n c r e a s e  p o p u l a t i o n growth substantially  is  adult  This mortality  mortality  reduced to  to a p o i n t  zero before  i m p a i r s moose p r o d u c t i v i t y  equilibrium).  However,  possibility  that  at  moose d e n s i t i e s  moose'knr )  wolf  p r e d a t i o n may d e c r e a s e  2  depensatory m o r t a l i t y  factor  al.  I  Therefore  1983).  wolf-moose stable  system  is  equilibrium  (higher  recruitment,  support the densities  hypothesis  by w o l f  equilibria  moose d e n s i t y If  in  if  wolf  least  In  level,  suggested. increase,  a single  1980).  i n an  Alternatively,  removal  to  will  rate)  a second h i g h e r  food c o m p e t i t i o n w i l l responses w i l l years).  In  be  test  the  =1.0  low multiple  2  be q u a n t i f i a b l e  this  paper,  the  equilibrium will  density  only  to  continues  results  by  these  in a long-term study  preliminary  its  be  equilibrium created Obviously,  the  animal•km" ).  t h e moose p o p u l a t i o n  indicated.  a  would  be s t o p p e d when  doubled ( i . e . ,  if  area  r e g u l a t e d and near  m a i n t a i n e d at  order  low d e n s i t y  the  and an upper  s u b s e q u e n t y e a r s t h e moose p o p u l a t i o n d e c l i n e s  former  to  to v e r i f y  et  t h e moose p o p u l a t i o n  moose a r e  predation.  has a t  as a  1981; Gasaway  p o p u l a t i o n growth  that  hypothesis,  in  a  >0.5  reduce wolves  naturally  An i n c r e a s e higher  to  of  the  1 9 7 6 : 2 3 6 ; Van B a l l e n b e r g h e  where t h e moose p o p u l a t i o n was equilibrium.  al.  c h a r a c t e r i z e d by a l o w e r  (Clark  where  presence  (i.e.,  and a c t  am a l s o a t t e m p t i n g  The e x p e r i m e n t a l d e s i g n was  stable  recognize  (Walters et  depress  food competition  (i.e.,  low d e n s i t y  higher  I  may  (>10  accumulated  87  during  the  first  programme i s  four  y e a r s of  the  ongoing wolf  reduction  presented.  STUDY AREA AND METHODS  The s t u d y  was c o n d u c t e d i n La V e r e n d r y e  southwestern  Quebec  (Fig.  14),  from June  The moose p o p u l a t i o n has been l i g h t l y 15 y e a r s  (3-4%  annually)  d e t e r m i n e d by n a t u r a l and C r e t e  2  bounded by a  900 km scat  (Fig.  2  forest  insure  10-km b u f f e r  14).  Wolf  km.  interviews. 1 July  al.  scats  a wolf  for  the  was  basically  1981a;  past  Messier  on m o t o r c y c l e s  site.  patrolling  registration  office  were of  for  a  and 6 weeks  1-week  l e n g t h of  on  I  recorded low  6.9  ± (SD)3.0  form a permanent  reserve  at  the  to  period.  48-km observers  20 km«h.  patrol.  interviewed the  8,  sufficiently  a maximum speed of  were c o l l e c t e d d u r i n g e a c h Moose h u n t e r s  10, 8,  counts  C o u n t s were made by two  at  of  scat  r o a d segments had an a v e r a g e  i n each study  700  Weekly  W i t h i n each a r e a ,  to  of  was d e t e r m i n e d by  remained n o t i c e a b l e  selected  area  status  and l a s t e d  1983.  removal  and a n o n - r e m o v a l a r e a  w h i c h human use was  They were r a n d o m l y  circuit  scats  for  zone,  population  from 1980 t o  roads  that  Suitable  density  ( C r e t e et  were s e l e c t e d :  r o a d s began on  respectively all  1980 t o M a r c h 1984.  harvested  stable  regulation  c o u n t s and h u n t e r  forest  its  reserve,  1984).  Two s t u d y a r e a s km  but  game  compulsory  from 15 September  t o 20  All  88  76° 00 W  76°45 W  1  T  LAVERENDRYE  \.  RESERVE!—-)  x  WOLVES KILLED: o Nov.'81- Feb. '82 * Nov. '82- Feb. '83 a Nov.'83-Feb. '84  .  r  V 10-km BUFFER ZONE  o  47 15 N  47 15' N  L WOLF-REMOVAL AREA  47 45' N  76° 45' W  Fig.  14.  Description  and l o c a t i o n s  76° 00' W  of the study area f o r the wolf  of the wolves  killed  removal  experiment,  b e t w e e n November 1981 and M a r c h 1 9 8 4 .  89  October.  One p e r s o n of  he o b s e r v e d d u r i n g h i s  each p a r t y 1-4 days  he h e a r d w o l v e s h o w l i n g . the  number of  c o n d u c t e d d u r i n g the  first  were r a n d o m l y  plot in  twelve  fixed-wing a i r c r a f t  =120 m).  r e p o r t e d the  number of  moose seen p e r p l o t addition, from t h e  (submitted)  coverage  to  the  calves  were  Additional  in  calves  plots  for  et  Number of  ratios,  and a n t l e r size  survival  d u r i n g the p a r t u r i t i o n cows  derived  al. 100 km f l o w n from  was  total  estimated  sex of  criteria,  (Roussel  adults  and  1975).  was g a i n e d  Females  In  bias.  From h e l i c o p t e r ,  by r e l a t i v e  Afterwards  visibility  calfrcow  from v u l v a - p a t c h  ground  aircraft,  were  moose seen p e r  in both study a r e a s .  born.  the  patrolled  above  f o u n d by h e l i c o p t e r  surveys.  i n f o r m a t i o n on c a l f  intensively  were  each  i n d e x of moose d e n s i t y . within  number of  terms of  identified  c o l l a r e d cows  of  estimates  density  during January a e r i a l ascertained  of  (6X10  2  Within  altitude  one on e a c h s i d e  was used as an  the  60 km  .  500 m i n t e r v a l s  c e n s u s e s and c o r r e c t e d  Recruitment,  was  at  study area  r e g r e s s i o n p r e s e n t e d by C r e t e  wherein  was c o r r e l a t e d  in each  of  surveys  each  moose s e e n t o a n a v i g a t o r .  moose d e n s i t y simple  h u n t e d and  January  Four p l o t s  ( s p e e d =165 km«h;  Two o b s e r v e r s ,  the a r e a  days  noted.  weeks of  1979).  10-km t r a n s e c t s  of  was d e t e r m i n e d by a e r i a l  three  selected  wolves  h u n t i n g , and how many  were a l s o  status  y e a r ( C r e t e and S t - H i l a i r e km)  of  The l o c a t i o n  hunting-days  Moose p o p u l a t i o n  was a s k e d how many  from  radio-  were m o n i t o r e d  p e r i o d to a s s e s s  the  number  were l o c a t e d a p p r o x i m a t e l y  at  90  1 month i n t e r v a l s Wolf  to determine  removal  experiment.  began d u r i n g t h e  From November  c a p t u r e d by l o c a l aerial Also,  trappers  shooting sessions wolves  were k i l l e d  removal area  to  calf  the  losses. third  year  of  the  to March,  wolves  or  from h e l i c o p t e r .  the  killed aim was  when t h e y  10-km b u f f e r  to  were  either  remove e n t i r e  were t r a c k e d  from  During packs. the  zone.  RESULTS  From November were k i l l e d zone. and  1981 t o F e b r u a r y  in  the  The a n n u a l  10 i n  (n=20),  700-km  removals  1983-84.  there  2  For  removal area were:  of  helicopter  was n e c e s s a r y  trapping effort  for  people  is  males,  4 females,  never  totally  females —  were  Weekly  population Whitney the  unknown.  removal  the Pups  remaining  in  7 in  the  An a v e r a g e  wolf  killed  13 w o l v e s  Thus,  a l t h o u g h at  1982-83,  buffer that  of  zone, they  6.7 h of  (n=l8).  The  c a p t u r e d by  local  (n=17;  reproductive least  wolves  10-km b u f f e r  were c a p t u r e d e a c h y e a r  1 unknown).  eliminated  1981-82,  31  observations  area. per  and t h e  killed  from t r a c k  were u s i n g p a r t time  14 i n  most w o l v e s  was e v i d e n c e the  1984, a minimum of  12  adults  were  8 m a l e s and 5  killed. scat  in  the  counts  (Table  18)  indicated  removal area  was  successfully  U-test[20,14]  non-removal area  = 237.5,  P<0.01)  but  that  the  wolf  reduced  (Mann-  remained s t a b l e  (U-test[20,14 ] = 153.5,  P>0.10).  in  Hunter  91  Table  18.  Indices  i n the removal (1980,  1981)  area  of wolf  populations  (R) b e f o r e  i n the non-removal  initiation  and d u r i n g t h e programme  No.  scats  per  100 km • week  of the wolf  (1982,  No. w o l v e s 100  reduction  (N-R)  and  programme  1983).  seen  per  hunting-days  (No. of  area  sightings)  No. days w i t h per  100  howling  hunting-days  (No. days  howling)  N-R  R  8.3(2)  7.0(6)  20.8(5)  2.9(3)  2.1(2)  12.4(13)  10.3(10)  5.0(7)  1.0(1)  3.6(5)  8.1(8)  N-R  N-R  R  1980  5.6  3.1  1981  4.8  13.1  3.4(3)  1982  6.0  2.1  1983  8.0  0.4  •  R  Pre-removal  Post-removal  92  interviews  gave  inconsistent  i n d i c a t e d by t h e percentage Small  scale  densities  the wolf  vs  0.34  year  testflO]  from t h e  18).  independence analysis  interviews  of  these  when u s e d on a  but  wolf  0.28 a n i m a l « k m ~  non-removal area  2  in the  (Table  removal  19).  After  r e d u c t i o n , t h e moose d e n s i t y  i n c r e a s e d t o an e s t i m a t e d 0.73 a n i m a l - k m " (t-test[l0]  remained s t a b l e  = 0.38,  P>0.10).  = 2.38, in  the  P<0.05; data  of  1981+'82  non-removal a r e a  (t-  U n u s u a l d e e p snow c o n d i t i o n s moose d e n s i t y  for  the  in  2  in  second year  t h e programme. B e f o r e wolf  approximately one y e a r the  not  (Table  statistical  of h u n t e r  1984 p r e v e n t e d a s s e s s m e n t of of  of  was  r e d u c t i o n programme (1981 +'82 ) , moose  i n the  of  removal area 1983)  s i g h t i n g s , but  possible lack  were e s t i m a t e d a t  significantly the  reduction  s h o u l d be r e c o g n i z e d .  and a t  first  a wolf  hunters heard wolves  The l i m i t a t i o n  Before  the  days t h a t  wolf  o b s e r v a t i o n s o b s c u r e any  results.  area,  f r e q u e n c y of  sample s i z e s and t h e  between  small  of  results;  of  treatment,  calf:cow  two y e a r s o f  cows were c o u n t e d i n t h e  A higher  calf:cow  calf  survival  the  lower in  (Table  the  improved s u r v i v a l  ratios 20).  increased to  39:100 the  were After  68:100  subsequent  in  year.  programme, 52 c a l v e s : 1 0 0  removal area  (significantly  i n d i c a t e d by the  ratio  but d e c l i n e d t o  first  removal a r e a  (l98l+'82),  40:100 i n both study a r e a s  removal area  Summing t h e  reduction  at  and 3 6 : 1 0 0 i n P=0.07, T a b l e  removal a r e a of  calves  was  t h e non20). also  associated  with  93  Table 19. methods)  Average  (SE)  and e s t i m a t e s  and i n t h e r e m o v a l  number o f moose o b s e r v e d p e r 6 0 - k m (SE)  area.  o f t h e moose d e n s i t y  Results  in  in the  plot  (see  non-removal  1983 w e r e d u r i n g t h e w o l f  reduction  programme.  N o . moose  Moose  observed 2  p e r 60-km  density  No./km  plot  2  Non-removal  Removal  Non-removal  Removal  area  area  area  area  Pre-removal 1981  4.7  (0.8)  2.8  (2.7)  0.36  (0.12)  0.23  (0.21)  1982  4.3  (0.6)  4.3  (2.3)  0.32  (0.11)  0.32  (0.18)  5.0  (1.5)  10.7  (2.7)  0.37  (0.14)  0.73  (0.22)  Post-removal 1983  94  Table  20.  Calf-cow  area before  ratios  initiation  of the wolf  and d u r i n g t h e programme two-tailed  Z-test,  i n the non-removal  (1983,  reduction  1984).  a r e a and i n t h e programme  The r a t i o s  (1981,  removal 1982),  were compared w i t h  a  = = 0.10.  No. c a l v e s :  Non-removal  area  100 cows ±90% CL  (n)  Removal  area  Z-test  Pre-removal 1981  65 ± 30  (43)  30 ± 1 8  (35)  n.s.  1982  24±11  (56)  52 ± 2 5  (38)  n.s.  1983  45 ± 1 7  (61)  68 ± 2 9  (47)  n.s.  1984  30 ±11  (74)  39 ± 1 8  (50)  n.s.  1981 + 8 2  39 ±11  (116)  40 ± 15 ( 7 3 )  1983 + 8 4  36 ± 8 ( 1 5 4 )  52 ± 16 ( 9 7 )  Post-removal  n.s. P=0.07  95  radio-collared survival (Fig.  rate  females. of  In  the  removal  area,  =65% compared t o =50% i n  calves  the  had a  non-removal  area  15).  DISCUSSION  A l t h o u g h the be  results  improved, they  are  predation maintains the  food c a r r y i n g The w o l f  level. 2  per of  100 km the  the  1981-82,  density  Keith  (from  (1983)  densities  area  and i t s  1982-83,  before  averaged  wolf 1.4  number of  Moose d e n s i t y  0.3,  well  below  et  wolf  animals*100  km"  in  (1984b)  the  wolf  As an  estimate  density from  2  first  (0.28  0.73 m o o s e « k m ~ ) ,  indicated.  in press  year  the  is  1980-81  a  to  a).  removal Scat  population.  I  a trend  the  100 h u n t i n g - d a y s  However, to  in  p o p u l a t i o n to d e c l i n e .  i n c r e a s e d d u r i n g the  increment  (i.e.,  argued that  w o l v e s seen p e r  at  animal  1983-84.  removal,  al.  likely  and 0 . 5  r e d u c t i o n programme.  nonetheless  that  zone  radio-tracking; Messier  and P e t e r s o n  suggested a reduction  was  buffer  0.7,  and  r a t e e x c e e d i n g 30% c a u s e s a w o l f  density  can  and s u b s e q u e n t l y h e l d  represented  intensive  c o u n t s and t h e  hypothesis  removal area  1981-82,  removal  in  non-removal area  in the  40% i n  removal  2  w i t h the  precision  capacity.  wolf  wolf  1983-84  consistent  and t h e i r  moose p o p u l a t i o n s at  least In  2150 k m ) ,  preliminary  population  r e d u c e d by a t this  are  of  the  have some r e s e r v a t i o n s 2  Aerial  survey  wolf  about  but precision  100 r  a)  ©  REMOVAL  (—•—)  80 -  ®  w  53  CD > 60 CO  E]  ©  ©  0  EH Ea  O  NON-REMOVAL  "CO 40 >  (—o—)  CO 20  0  L  I  M  J  L  J  A  S  O  N  O  J  J  F  I  M  1  A  MONTH  Fig.  15.  Percent  survival  f e m a l e moose i n t h e w o l f  of calves  removal  associated with  and t h e n o n - r e m o v a l  radio-collared area.  Total  n u m b e r s o f c a l v e s f o l l o w e d t h r o u g h t h e s u b s e q u e n t month a r e i n and  squares.  circl  97  must be i m p r o v e d t o q u a n t i f y increment. t o meet  Moose i n v e n t o r y  this  Higher ratio,  (Crete et  recruitment,  represents  population al.  goal  1983).  calves:100  In  this  cows t o  and d e c l i n e d t o second year yearling  1983, y e a r l i n g m a l e s  of  all  males  in the  non-removal area assume s i m i l a r  percentages  for  interpretation  the  of  the  non-reproductive  (i.e., higher  cows  ^1.5  recruitment  23% (n=31)  in  P<0.05).  females,  the  I  number of  indication of  the  If  removal area  was an  (n=17)  and 39 that  wolf  be s u p p l e m e n t e d i n t h e  estimates  where  sample  future  sizes  doubled. survival  cows t r a c k e d these  removal  Probably  = 2.13,  82 i n t h e  r a t e of c a l v e s  f e m a l e s gave a l e s s  of  et  r e p r e s e n t e d 53%  but o n l y  Thus t h e r e  recruitment  The e v a l u a t i o n  number of  of  yearling  o l d was  a moose  wolf  was much i m p r o v e d i n b o t h y e a r s  s h o u l d be a p p r o x i m a t e l y  small  of  cow p o p u l a t i o n  This conclusion w i l l  radio-collared  year  by a l a r g e r  Z-test  of  calfrcow  (Gasaway  second y e a r .  removal area  (two-tailed  1985  i n c r e a s e d from 40  (spiked antler)  non-removal a r e a .  by more p r e c i s e  predation  As a r e s u l t  cows ^ 2 . 5 y r  recruitment  the  6-month  indicator  first  r e d u c t i o n was c r e a t e d  i n our c o u n t s ) .  reduction.  the  the  in  submitted).  recruitment  39:100 f o r  in  the  al.  68:100 i n  old  the  be r e v i s e d  important  study,  yr  in  technique w i l l  from i n t e n s i v e  cohort entering  calves:100  population  as e x p r e s s e d by t h e  t h e most  relieved  more p r o p e r l y  pronounced response.  to date  preliminary  associated  necessitates results.  a  with The  cautious  98  Black density are  of  the  calf  experiment  site  the  reserve.  on t h e  Management  to  et  al.  removal  is  that  example,  true,  in c a l f  in  1983 w i t h i n  b e a r s as c a l f  They and  survival  even  A bear  a third  should provide  the  'natural  the d e n s i t y  interference ( C r e t e at  study  some  predators.  Alternatively,  al.  (Connolly  immediate p u r p o s e of determine  low d e n s i t i e s  the  if  low  a dilemma.  surplus'  With t h i s (e.g.,  and a v a i l a b l e  moose  by p r e d a t i o n .  face  harvestable  1981).  is  only  If They  limited. yield 0.2-0.3  alternative,  the  1983 p r o v i n c e  food resources  wide  could  moose p o p u l a t i o n s .  may a d v o c a t e  predator-moose-vegetation perspective,  The  possible  ungulates  i n Quebec was e s t i m a t e d a t  support higher they  t h e many  o f f e r i n g a maximum s u s t a i n e d  s u c c e s s was-8%),  be u s e d t o  one of  moose managers  hunting  this  increase  to e x p e r i m e n t a l l y  s u c c e s s would r e m a i n  the  1980),  1983).  hunting  not  al.  p o p u l a t i o n growth of  is  hypothesis  2  et  represents  this  moose«km"  (Franzmann  initiated  r e g u l a t e d at  without  1984).  r e d u c e d more s e v e r e l y .  p o p u l a t i o n s are  For  al.  i m p o r t a n c e of  stimulate  wolf  may a c c e p t  a minimum  et  This project  control  1978; Gasaway present  at  implications  Predator options  was  sites  (Lachapelle  to the  p o p u l a t i o n was  removal  insights  2  predators  impose a l i m i t  wolf  in  common i n b o t h s t u d y  0.25 a n i m a l ' k n r  potential  probably if  bears are  that  the  system are  and c o n s i d e r i n g t h e  pristine  nature  of  p e r t u r b e d by man.  large  demand f o r  moose  In  99  hunting policy  in may  populations  some a r e a s , include  an  options  by means of  integrated to  ungulate  simultaneously  trapping  and h u n t i n g .  management manage  predator  100  Chapter  V  GENERAL DISCUSSION AND CONCLUSIONS  Predator-moose-vegetation graphical  representation  a moose p o p u l a t i o n  is  (Fig.  in a s i n g l e ,  suggested  for  forage  ungulate  with  In  and a c c e n t u a t e s  growth  no l o n g e r  In  the  exists. of  In  of  this  first  depressive  effect  second v a r i a n t induced) This  type  growth at  is  but  (i.e.,  a density  detrimental l e s s e n i n g of  effect  (D„),  1976;  as  Sinclair  until  for  restrains  population  s y s t e m s may have  predation  competition  for  the  increment  to create  forage  an  having a The  3  implies  that  where f o r a g e  predation  reduces  outcomes  food-induced e q u i l i b r i u m , D ) .  (food-induced)  (D,),  possible  sufficiently  a system c h a r a c t e r i z e d  system  growth,  competition  a c o n t i n u u m of  variant,  without  and an upper of  (Caughley  chronic mortality  a moose p o p u l a t i o n , but n e v e r  equilibrium density  equilibrium  situation,  predators,  3 variants,  the  species  logistic  area,  possible.  presence  qualitatively  a predator-free  increasing population density  fecundity is  In  upper d e n s i t y  1977; M c C u l l o u g h 1 9 7 9 ) . nutritious  16).  expected to d i s p l a y  resulting  other  d y n a m i c s c a n be summarized by a  but a t  by a l o w e r  equilibrium  predators  higher  rate permits  density.  can s t o p  c o m p e t i t i o n has  (predator-  population  no  moose d e n s i t i e s ,  p o p u l a t i o n growth  a until  101  Fig.  16.  The p r o p o s e d c o n c e p t u a l  dynamics.  In p r e d a t o r - f r e e  to s t a b i l i z e 3 variants (Dg),  2)  of  at  predation  predator-moose-food  moose p o p u l a t i o n s  are p o s s i b l e :  predator-induced  is  of  (D^ e q u i l i b r i u m ) .  system behavior  induced e q u i l i b r i u m model,  areas,  high density  a lower,  model  and 3) envisioned  by an i n v e r s e d e n s i t y - d e p e n d e n t  1)  equilibrium  a single,  are  hypothesized  Where p r e d a t i o n a single, (D ) n  supply  upper  occurs, equilibrium  and an u p p e r ,  food-  lower e q u i l i b r i u m  (D-|).  In  as h a v i n g a d e n s i t y - d e p e n d e n t  phase  followed  phase  (inset).  this  102  competition third  for  variant  density  (D,)  forage  consists  effect  population  growth at  of  the  (D ).  predation.  In  this  a  low  case,  the  a high predation  rate  higher  (negative  densities  The  2  a system s t a b i l i z i n g at  element  precludes  i n d e t e r m i n i n g the  r e l a t i o n s h i p between  Walters et  al.  the  of  impact  population  increment,  (1981), wolf  predation  and Gasaway  predation  decline.  This  et  system's  behavior  r a t e and moose  density.  al.  increases  s o - c a l l e d depensatory  a moose p o p u l a t i o n t o e x t i n c t i o n ,  never  reported  in  the  literature.  mechanisms must be i n v o l v e d .  In  this  lower  dependent phase (inset  of  Fig.  to  systems). predation  16).  Holling  because  1978 f o r  At v e r y  they  moose d e n s i t y  influence  (1959)  examples  in his  s t r e s s e d and t h e r e f o r e  above  the  feedback  have  of  wolf  predation  the  predation  density-  relationship  (see  predator-prey moose•knr  because wolves  r a r e or a b s e n t ,  density  phase  work  (<0.2  t e n d t o consume a l t e r n a t i v e  was  pioneering  in arthropod  negligible  some  density-dependent  low moose d e n s i t i e s  increases  I  Overall,  found a s i m i l a r  by w o l v e s a p p e a r s  nutritionally  study,  inverse  that  s u c h an e v e n t  be c h a r a c t e r i z e d by a  f o l l o w e d by an  w i t h mammalian p r e d a t o r s also Hassell  but  range of moose d e n s i t i e s .  r a t e by w o l v e s a p p e a r s  argued  i m p a c t may  Consequently,  demonstrated the d e n s i t y - d e p e n d e n t the  (1983)  d u r i n g a moose  force  at  effect  16). The c r u c i a l  is  of  due t o heavy  persistent  Fig.  has a r e g u l a t o r y  2  ),  are and  food r e s o u r c e s . t h r e s h o l d of  also When  =0.2  103  moose-km ,  wolves can c o l o n i z e the  - 2  initial  struggle  territories  to  intensively  c o l o n i z a t i o n phase moose«km" )  providing a strong  must e x p l o i t  (Messier  in press a ) .  deferred  2 reasons:  reproduction)  ( P a c k a r d and Mech  to  impact  1)  social  of  et  s u b s t a n d a r d moose i n d i v i d u a l s  are  compose a g r e a t e r  wolf's  al.  fraction  and  wolf  0.5  dramatically, higher  predation  likely  (territoriality, population  1983),  and  more a v a i l a b l e kills  this  growth  2) and  would  (Peterson  et  1984b). The r e a l i z a t i o n  between  annual  increment  limited  curve  wildlife  or  curve  (1976).  w i t h a range  of  nil  in  may n o t  Rather, the  r a t e peaks  densities (Fig.  16).  at  at  c o n s e q u e n c e s of  the  general of  which  initial  r e s p o n s e of  important For  example,  shape as  f o r m of  the  suggested  the  annual  p r e d a t i o n may be b i m o d a l , the h a r v e s t a b l e  such a " p r e d a t o r  surplus  decline  wolf  pit"  increases  when h u m a n - e x p l o i t a t i o n  h i g h moose d e n s i t i e s . an  somewhere  is  T h i s phenomenon p o s e s a p r o b l e m  an u n e x p e c t e d d e c l i n e  lag  has  have a b e l l  presence  managers b e c a u s e  liberalized  typical  predation  on moose p o p u l a t i o n d y n a m i c s .  increment  by C a u g h l e y  of  that  low and m o d e r a t e moose d e n s i t i e s  implications  risk  of  During  At d e n s i t i e s  wolf  al.  their  their  0.2  behavior  would r e s t r a i n  1980; P a c k a r d  in  increase  feedback mechanism. 2  for  between  rate appears  0.5-1.0 moose-km" , the  decreases  but  they  (approximately  predation  2  than  subsist  system,  Moreover,  for the  is  the  can be a g g r a v a t e d  numbers t o d e c r e a s i n g  by  the  ungulate  1 04  densities and Page  (Mech and K a r n s  predator  little  pit  proposed at  is  information  hypothetical,  this  ungulate  time.  is  species.  This effect  stabilize  at  likely  predator.  In  to  al.  available,  1983;  Peterson  if  the  I  1983), of  the  such s i t u a t i o n s  other  pit  of  ungulate t h a t moose  predation.  of  predators  the p r e d a t o r will  impact  possibility  due t o w o l f  recruitment  the  with  the p r e d a t o r  the  be a b s o r b e d by t h e  expect  form of  where moose c o h a b i t  a second s p e c i e s  bear),  the  some p r e d i c t i o n s may be  Carbyn  decreases  moose p o p u l a t i o n s a r e densities,  but  areas  (e.g.,  low d e n s i t y  Alternatively,  deep because  is  pronounced because p a r t  predation  black  In  species  s h o u l d be l e s s  (e.g.,  et  1983).  Because  other  1977; Gasaway  pit  is  present  t o be  relatively  be r e d u c e d by t h i s  second  (as  in  southwestern  Quebec),  more l i k e l y  to  stabilize  low  and may n e v e r  increase  naturally  at  to  higher  densities. Gasaway (Slobodkin system.  et  al.  1974)  is  Evidence  p o p u l a t i o n s are  (1983)  fact  Alaska,  least  30-40 y e a r s or any (density  and C r e t e between  (1984)  1967 t o  that  some n a t u r a l l y  remarkably  Park,  winters  Haber  a "prudent"  r e q u i r e d to e x p l a i n a s t a b l e  exists  in  argued that  (1977)  stable.  = 0 . 3 - 0 . 4 moose•knr ) . 2  c o u l d not d e t e c t 1983.  Among t h e  In  Denali  National  irruptions  for  following  severe  In  Messier  area  any c h a n g e s 3 study  moose-wolf  r e g u l a t e d moose  r e p o r t e d no major  obvious crashes  predator  H,  i n moose  areas,  at  density  predation  rate  105  increased  sharply  moreover,  predation  j u d g i n g by t h e able  they  predators. range  rate  d i d not  killing  an  were v e r y  In  this  stable  to  r a t e and f o o d i n t a k e .  W o l v e s were  in  t h e moose p o p u l a t i o n rather  than  in predation  precisely  prudent  rate within  an e f f e c t i v e  predation  represents  (1981), the  a  narrow  feedback  predator-moose-vegetation  as p o s t u l a t e d by Peek  where  and  i t s maximum,  constitutes  sense,  19.3% ( a r e a H ) ,  appear  efficient,  a l w a y s dynamic  be f a i r l y  L)  t o be a t  increase  of moose d e n s i t i e s  not  (Area  The s h a r p change  mechanism. are  low  to prevent  because  from 6.1%  systems but may  regulatory  mechanism. Evidence  also  exists  that  p o p u l a t i o n s may u n d e r g o major b e i n g the From t h e  I s l e Royale  moose numbers a r o u n d t h e  competition (Blood  densities  mortality.  decreases  at  the  same t i m e  that  the  net  f e e d b a c k mechanism a t population density.  growth  the  (above).  result  D  and D .  2  of  it  winters  on  the  recruitment  predation these  is  in  forage  and p r e s u m a b l y  would be a weak  change  At  3  effect  h i g h moose d e n s i t i e s ;  severe  changes  Because  antagonistic,  r a t e does not  Occasional  impact  example  chaotic  - 2  However,  best  1983).  rather  1983)  moose  and Page  t o =4 m o o s e - k m ) ,  and H a a g e n r u d  p o p u l a t i o n processes are believe  predict  upper e q u i l i b r i a (1  the  (Peterson  induces a density-dependent  1973; S a e t h e r  chronic  I  regulated  fluctuations,  population  conceptual model,  h i g h r a n g e of  naturally  on  likely  2  reasonable  to  regulatory that  rapidly  may c a u s e  is,  with  population  important  1 06  fluctuations 1977).  It  Royale  over  essence  of  may w e l l  disturbances predation;  the  15 y e a r s  these  the  lack  stochastic  of  the  explanations  al.  et  1983; P e t e r s o n  (1977)  diverts  e q u i l i b r i u m than lower  will  predators  this  depressive  effect.  from D„  create  Quebec  3  stressed,  an u p s u r g e of  indication  that  (Messier  forage  (Fig.  3  these  proposed  Both  16).  population  are  enrichment (Peek  of  two p r e d i c t i o n s a r e  that  that  the  true  in  be  must  be  will  There  Alternatively,  density.  conclude  to  forage  1980).  density-dependent I  how much  argued  moose i n d i v i d u a l s  regarding D,:  true,  lower  Two p r e d i c t i o n s can  and 2)  this  that  from  referring  1)  1984).  But,  authors  p r e d i c t i o n can be f o r m u l a t e d  appears  et  c o m p e t i t i o n has a  they  moose d e n s i t y  up t o  remains  (Bergerud  situation.  or D „ :  and C r e t e  high  view  p o p u l a t i o n t o a new and  Therefore,  to D  of  phenomenon  plausible  an u n g u l a t e  where  regarding D  nutritionally  are  new e q u i l i b r i u m be?  to a l e v e l  cyclic)  Although t h i s  basic  in  1984a).  an u n g u l a t e  prevent  to  was  f e e d b a c k mechanism a t  in a p r e d a t o r - f r e e  cannot  formulated  al.  Isle  1983)  effect  and M c C u l l o u g h (1979)  increasing  decrease  and Page  contrast  reason.  u n t e s t e d and o t h e r  predation  (in  an e f f e c t i v e  fundamental  (Peterson  phenomenon o b s e r v e d on  (Peterson  my i n t e r p r e t a t i o n ,  Caughley  high densities  e x a g g e r a t e d by t h e m u l t i p l i e r  b e i n g the  represents  be t h a t  the past  one of  density  moose numbers a t  is  southwestern one  p r e d a t i o n must  Because  no  this  moose p o p u l a t i o n s a r e  be  prediction either  107  predator-regulated,  or  predator  multiple  equilibria  system).  currently  attempting  to  pit  (i.e.,  Messier  (1984)  between  these  are  s t a b i l i z e d at  2 alternatives  low d e n s i t y  by means of  below  a  Crete  and  discriminate  a wolf  removal  experiment. At  this  time,  could s t a b i l i z e reduction density  at  it  is  high d e n s i t i e s  that after  programme; moose may r e t u r n  due t o heavy  population.  predation  Furthermore,  equilibrium  remains  utilization  and n a t u r a l  outcomes.  uncertain  to  the  after  to  be d e m o n s t r a t e d , still  population  a temporary their  recovery  stability  crashes  a moose  of  wolf  former of  the  a high  habitat  represent  low wolf  density  overpossible  108  LITERATURE  CITED  A d d i s o n RB, W i l l i a m s o n J C , S a u n d e r s BP, F r a s e r D (1980) R a d i o t r a c k i n g of moose i n t h e b o r e a l f o r e s t of n o r t h w e s t e r n O n t a r i o . Can. F i e l d - N a t . 94: 269-276. A d o r j a n A S , K o l e n o s k y GB (1969) A manual f o r t h e i d e n t i f i c a t i o n of h a i r s of s e l e c t e d O n t a r i o mammals. O n t . D e p t . L a n d s F o r . , R e s . Rep. W i l d l . N o . 9 0 . 47 p p . 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Manage. 4 0 :  W a l t e r s C J , S t o c k e r M, Haber GC (1981) S i m u l a t i o n and o p t i m i z a t i o n models f o r a w o l f - u n g u l a t e s y s t e m . I n : F o w l e r CW, S m i t h TD ( e d s ) D y n a m i c s of l a r g e mammal p o p u l a t i o n s . John W i l e y and S o n s , New Y o r k . p p . 3 1 7 - 3 3 7 . Zimen E (1976) On t h e r e g u l a t i o n of Z. T i e r p s y c h o l . 4 0 : 3 0 0 - 3 4 1 .  pack  Zimen E (1982) A w o l f pack s o c i o g r a m . I n : Paquet PC (eds) W o l v e s of t h e w o r l d : b e h a v i o r , e c o l o g y , and c o n s e r v a t i o n . (New J e r s e y ) , p p . 2 8 2 - 3 2 2 .  size  in  wolves.  H a r r i n g t o n FH, p e r s p e c t i v e s of Noyes, Park Ridge  1 17  Appendix  SOLITARY  LIVING AND EXTRA-TERRITORIAL MOVEMENTS OF WOLVES  IN  RELATION  TO SOCIAL  STATUS AND PREY ABUNDANCE  INTRODUCTION  In  social  may p l a y  b i r d s and mammals, a central  regulation  role  attracted  sociality 1982;  In  space  predators  Bekoff  (habitat over  carcasses, (reviews  et  al.  factors  to  temporarily  in  the  the  decade.  1977;  within  The  Emlen  from i t s  2)  a framework 1) the  of  the competition  competition with  other  needed t o  rear  1981; M a c d o n a l d and  1982; M a c d o n a l d 1983;  propose t h a t , the  by:  assistance  i n Lamprecht  I  philopatry,  1983).  3)  and 4)  last  1977; B e k o f f  food r e s o u r c e s ,  1 and 2 g o v e r n apart  the  natal  and  s h o u l d h e l p t o comprehend  and B a r r e t t e  in p r e s s ) .  or of  operates  saturation),  1982; M e s s i e r  g r o u p members  The b e h a v i o u r a l  determined l a r g e l y  and abundance of  lupus), live  group l i v i n g  constraints  young o p t i m a l l y Moehlman  interest  1983; Waser and J o n e s  cost-benefit  for  dispersal,  1974; A r m i t a g e  Carnivora,  dispersion  1983).  such a n t e c e d e n t s  (Barash  Brown  of  biologists'  u n d e r s t a n d i n g of  of  i n g r o u p d y n a m i c s and p o p u l a t i o n  (Brown 1983; M a c d o n a l d  e c o l o g i c a l antecedents have  the d i s p e r s a l  in wolves  tendency pack  of  an  (solitary  (Cards individual living),  and  1 18  t o engage of  this  in  solo e x t r a - t e r r i t o r i a l  proposal, I  with t e r r i t o r y dispersal Packard  vacancy  strategy  and Mech  Here  I  of  that  ultimately  interpreted  age of  the  animals,  their  packs.  Observations  al.  Mech  of  rules  also  sex,  in  and  for  the  Zimen 1976;  research  to prey  spatial  field  abundance,  distribution  included  and Mech ( 1 9 8 1 ) ,  Van B a l l e n b e r g h e  published related  conjunction  extra-territorial  relation  and the  Fritts  extension  1983).  pack e x c u r s i o n s a r e  (1977),  (1982),  (1984)  (see  in  54 w o l v e s s t u d i e d d u r i n g a 4-year  Data are  and James  the  living  programme.  comparison.  set  pack members  on s o l i t a r y  As an  food resources  1980; M a c d o n a l d  report  movements o f  suggest  movements.  (1983),  for  Stephenson  and P e t e r s o n  observations  of  which  et  are  discussed.  STUDY AREA .AND METHODS  Wolf of  spatial  distribution  southwestern  February  reserve  1984. T h i s a r e a  Fifty-four tracked  2  2  by a i r p l a n e s .  a c c u m u l a t e d d u r i n g the  area  low p r e y  same s t u d y wolves  (47°N,  area  (HP) area  i n a 6400-km  77°W)  was d e l i b e r a t e l y  high prey  and a 3400-km (i.e.,  investigated  Q u e b e c , Canada  c o v e r e d a 3000-km reserve,  was  from June  selected  i n La (LP)  2000 h of  time.  the  II).  locations  flying  1980 t o  Verendrye  from 14 c o n t i g u o u s p a c k s were Some 4700 w o l f  area  because  s o u t h of  than Chapter  2  radio-  were W o l v e s were  it  119  visually rarely  observed  in  other  group s i z e ,  in  periods.  whether  open h a b i t a t ) ,  75% of  locations  Each wolf  or not  and t h e  the  all  during winter,  observation  the a n i m a l s  identity  of marked  but  included  were seen  the  (i.e.,  individuals.  C a p t u r e d w o l v e s were c l a s s i f i e d as e i t h e r  pup (<12 m o n t h s ,  starting  1 May),  or a d u l t  months).  Pups were aged a c c u r a t e l y  patterns. or  Yearlings  aged a f t e r  canine  that  reproductive  to  or  t o o t h wear,  if  so,  from t o o t h  of  of  known age  living  30 A p r i l  for  from the a i r  Frequency  (winter  period)  (n=1l92)  abundance  territory.  of  prey  p o r t i o n of Data  I  In  this  was c e r t a i n  on t h e  full  set  of  (HP  the  age of  or L P ) .  these  I  tables  (i.e.,  animals,  from known l o n e w o l v e s ,  or  not  at  and Gwhich  distant their  used s i m i l a r  observations  I  pack  1981: 7 4 7 - 7 6 4 ) . F r e q u e n c i e s  to  paper,  reliable  were b a s e d on c o n t i n g e n c y  from t h e p a c k s  from 1  whether  was a c c o m p a n i e d by o t h e r  analyses  500 m) were r e l a t e d  the  believed  when w o l v e s  ( S o k a l and R o h l f  relate  is  The  each year  w o l v e s were o b s e r v e d a p a r t  the  status.  their  only  observations  observations  than  from  pups  c o u l d be s t u d i e d r e l i a b l y  which  tests  as  would have been y o u n g , non-  used o n l y  members.  (n=3)  and r e p r o d u c t i v e  from a i r p l a n e s .  seen  (trapped  assumed age  c o u l d be o b s e r v e d d i r e c t l y  a wolf  eruption  were m i s t a k e n as a d u l t s  they  (>24  adults.  Solitary December  n=5),  yearlings  be m i n i m a l ;  (12-24 m o n t h s ) ,  were e i t h e r  death,  lengths,  possibity to  yearling  in  outside including  sex,  analyses the  more and to  pack's  animals  1 20  during their eliminated because I travel the  actual  or assumed d e f i n i t i v e  from t h e s e  they  analyses  represent  and were t r e a t e d  a different  defined e x t r a - t e r r i t o r i a l 5 km beyond t h e  pack.  Such a r e a s  containing member(s)  95% of  30 A p r i l ) .  In  the  the y e a r - l o n g  3 cases  by t h e  locations  o b t a i n e d d u r i n g the  current  where t h e  of  1-year p e r i o d  annual  b o u n d a r y of  the  subsequent y e a r .  line  most d i s t a n t  distance wolf  of  insufficient  to  the  closest  territory.  The a p p r o x i m a t e d u r a t i o n  calculated  from t h e m i d - d a t e  the  territory  mid-date first not  and t h e  between  location  the  independent  last  fixes  took  the  within  of  of  last  the  the  t h e home was  location  inside  e x c u r s i o n , to  e x c u r s i o n and  territory. not  of  Elapsed  time  the  the was  bounded by a p r e - and a  5 days,  or  if  an a c t u a l  d u r i n g an e x c u r s i o n were assumed t o  b e c a u s e most days  (30-40  Therefore,  wolves had,  foraying  the  The minimum  point  the  not  or  occurred.  travelling  continue  I  to  could  each e x c u r s i o n  the  location  location to  of  between  e x c u r s i o n was  location  assumed d i s p e r s a l Radio  first  on r e t u r n  computed i f  post-excursion  the  data,  pack  an e x c u r s i o n was measured from  location  of  (1 May  territory  of  straight  territory  instrumented  d e f i n e d because or  any  convex-polygon  be p r o p e r l y  previous  separately  e x c u r s i o n s as b e i n g  b o u n d a r y of  closest  were  phenomenon.  were d e l i n e a t e d  the  dispersal,  or  locations  km)  were w i t h i n  from t h e  i n most d a y s ,  t o be  in  their  pack's the  be  1 or 2  territory.  opportunity  home t e r r i t o r y  either  by t h e  to  next  121  day.  On a v e r a g e ,  that  met my c o n s t r a i n t s  separated  Radio the  Because  1984, pack  data  of  set  fairly  in  this  locations  locations  of  solitary  interstices.  of  territory  between  Locations  the  whole  outside  1981)  were v i o l a t e d .  significant.  the  areas  were  area  statistical  All-tests greater  the  than  of  pack  from 1980 1980-84 were  formed d u r i n g  the  wolves.  fixes  Radio  buffer  zones,  was c o n d u c t e d  areas  minus t h e  or  as 2-km wide  interstices  bordering  territory  area  equalled  cores.  were e x c l u d e d .  tests  (Siegel  a s s u m p t i o n s of  except  relation  territories  each t e r r i t o r y ;  plus  in  from t h e  m o s a i c . The t o t a l  areas,  total  were u s e d i f  probabilities  also  the c a s e s  p l u s a 2-km s t r i p  territory  plus buffer  cores,  = territory  territories  Nonparametric Rohlf  cores  these  zones b o r d e r i n g i n t e r n a l l y  interstices,  animals  were a v a i l a b l e  most y e a r - l o n g  territory  externally  of  animals  extra-territorial  2 new t e r r i t o r i e s  Planimetry  In  were d e l i n e a t e d  analysis;  few  during  were c a t e g o r i z e d w i t h i n  = areas  fixes  territory,  territories.  such o b s e r v a t i o n s  and t h e  study covered  buffer  the  solitary  w o l v e s were a n a l y s e d  pack  territories  stable  follows:  Radio  but w i t h i n  solitary  used only  movements.  4 days.  of  independent.  fixes  I  each  pack,  c o n f i g u r a t i o n of  members,  to  r e c o r d e d an o b s e r v a t i o n  from t h e i r  assumed t o be  to  I  F-test  1956; S o k a l parametric  were t w o - t a i l e d ,  0.05 were j u d g e d t o be  not  and  tests and  122  RESULTS  Solitary  living  W o l v e s w h i c h were c o n s t a n t l y in  the  study  females) others  Out of  c o u l d not  pack o r i g i n .  territories  after  one pack  suggests'a (perhaps  710 km  natal  be d e s c r i b e d as because  territory  previous  the  they  area  regularly  area  (85% of  but  members t e m p o r a r i l y  to  from t h e i r  d u r i n g p e r i o d s e x t e n d i n g from a few d a y s  I  rejected  the  null  abundance and age yearlings  to break  hypothesis  (G[2]=18.48, ties  on t h e of  mostly  which  areas.  can  orientation In  contrast  increased  implies  that  visited.  observed s o l i t a r y  analysis  during  2 lone wolves  This observation  were b e i n g  across  2  association  w i t h a random familiar  doubt  located  n=93),  and g r o u p these  and 3  postcapture.  each lone a n i m a l c o v e r e d  dissociated  table  was  locations,  not  2310 km  territories  animal  Movements of  locations.  frequently  the c o n t i n g e n c y  (both  which c a s t s  13 months  3 pack  The l a t t e r  returned  new a r e a s  The most  across  2  "itinerant"  number of  occasionally  capture  familiarity  pack).  t o pack members, t h e w i t h the  2 animals  One l o n e a n i m a l c o v e r e d  9 months p o s t c a p t u r e .  within  their  d u r i n g a p e r i o d of  The s e c o n d c o v e r e d the  54 w o l v e s t r a c k e d ,  were uncommon  be a s s o c i a t e d w i t h e s t a b l i s h e d p a c k s ,  dispersed shortly  on t h e i r 7 pack  area.  observed alone  w o l v e s were pack social  groups  t o a few m o n t h s .  full  set  of  winter  i n d e p e n d e n c y between  From data,  prey  P<0.01).  A low p r e y  base  caused  more f r e q u e n t l y  with their  pack  (45%  123  of  locations  pups  i n HP v s  (26% vs  and 3 2 4 ) .  Because  of  this  t o a 2-way a n a l y s i s  separate  1).  solitary  age  living  I  as compared t o  (17% vs  prey  28% of  the  b a s e and  in  among pups was  (P>0.15),  1),  and sex of  but a q u a n t i t a t i v e  (G[1]=5.82,  observations  (Table  1)  i n d e p e n d e n t of the a n i m a l s  indicate  I  difference  both prey  (P>0.20).  (Fig.  1),  alone  P<0.01),  abundance of  prey  and sex s t r o n g l y  yearlings  leave  to  yearlings  area.  In  than y e a r l i n g between their  prey  effects  their  or pups  pack  addition, yearling (Fig.  1).  a b u n d a n c e and sex (Table  (Fig.  significantly  1).  57% of  to  wolves  winter  frequency  (28%;  G[1],  than  (Table  Both  tendency 1).  pack a f f i l i a t i o n  Male in the  l i v e d more  The n o n - s i g n i f i c a n t indicates  adults  P<0.01).  i n f l u e n c e d the  females  of and low  solitarily  interaction  an a d d i t i v i t y  of  1).  A d u l t s m a i n t a i n e d the c l o s e s t groups  terms  similarity  from a d u l t  temporarily  e x h i b i t e d a lower  males  in  a much g r e a t e r  (21%; G[1]=84.72,  prey  In  r e p r e s e n t e d t h e most l o o s e l y a s s o c i a t e d pack  observed y e a r l i n g s  observations  female  that  P=0.02).  Yearlings members.  sex)  classes.  pack d i s s o c i a t i o n , pups showed a q u a l i t a t i v e  (Fig.  2 6 % , n=390  p r o c e e d e d from a 3-  (i.e.,  The n o n - s i g n i f i c a n t G - t e s t s  abundance of  or a d u l t s  interaction,  Pups were o b s e r v e d a l o n e (Fig.  n=122 and 6 1 ) ,  3 6 % , n=242 and 53)  way a n a l y s i s within  80% i n L P ,  Nonetheless,  contact  abundance of  influenced solitary  living  of  with t h e i r  social  p r e y and sex adults  (Table  1).  (5) 85  (3) 61  80  70 (8)  60  183 (5) 122  50 40  (5) 53  30  (7) (7) 157 138  (17) 324  (9) 242  (3) 98  (12) 390  20  (14) 288  (14) 295  (29) 714  (15) 426  10  LP HP  PUP  F i g . 1.  LP HP  YRLG  LP HP  ADULT  Cf  9  PUP  c? 9  d" 9  Y R L G ADULT  p  Y  A  OVERALL  Percentage of sightings when wolves were observed alone and  separated from their packs in relation to prey abundance (LP or HP), age of the animals, and their sex.  Number of sightings is given above  the columns, and the figure in parentheses denotes the number of animals involved.  The portion of sightings during e x t r a - t e r r i t o r i a l  is s t i p p l e d .  movements  125  Table  1.  Summary o f t h e c o n t i n g e n c y  cl asses,  where t h e e f f e c t s  related to the frequency and t h e p o r t i o n The G - t e s t  that  prey base a wolf  levels  ( d . f . =1 f o r ( ).  all)  analyses,  (HP o r L P )  left  of these odservations  values  probability  of  table  and s e x  temporally  its  outside the pack's  are presented,  Observations  within  age were  pack  (A),  territory  as w e l l  as  (B).  their  w e r e c o l l e c t e d f r o m 1 December  to  30 A p r i l . Hypothesis  Prey  tested  base  Pup  Yearling  Adult  1.82  (>0.15)  21.86  (<0.01)  9.72  (<0.01)  Sex  0.52  (>0.20)  44.54  (<0.01)  22.42  (<0.01)  Interaction  1.76  (>0.15)  1.67  (>0.15)  1.50  (>0.20)  Prey  0.04  (>0.20)  8.30  (<0.01)  2.80  (=0.09)  Sex  0.02  (>0.20)  58.22  (<0.0T)  0.22  (>0.20)  Interaction  1.80  (>0.15)  2.52  (>0.10)  0.02  (>0.20)  base  1 26  As w i t h y e a r l i n g s , frequently the  than m a l e s .  frequency  their  adult  pack.  at  females  travelled  A low p r e y  which a d u l t s  Once a g a i n ,  the  base and sex s u g g e s t s t h a t  of  lack their  base  alone  more  significantly  increased  both sexes t e m p o r a r i l y  left  of  prey  interaction  effects  between  were a d d i t i v e  (Table  D. As w i n t e r increased P<0.01),  i n pups  usually  a wolf  (G[3]=9.47,  and a d u l t s  From e a r l y was  p r o g r e s s e d , the  May t o  the  yearlings In  predictable  sites  summer, a d u l t s u c h as  territory  activity  sites  r e c o r d e d 56 i n s t a n c e s  represented:  renew  of  not  pack  to  pattern, than  and w i t h no a p p a r e n t  extra-territorial  d i d not  or  it  areas.  they adults  during prolonged periods,presumably  from t h e p a c k ,  animals  whether  d e n s and pup r e s t i n g  more e x t e n s i v e l y  engaged i n e x t r a - t e r r i t o r i a l the  period),  independent  demonstrated t h i s  the  I  (summer  movements c o n v e r g e d  natal  throughout  Individual  (G[3]=16.7,  2).  with c e r t a i n t y  a p p e a r e d t o be more  t e n d e d t o wander  focal  dissociation  yearlings  end of November  definitely  the  pack  pack members. From m o n i t o r i n g movements  A l t h o u g h some y e a r l i n g s  dissociated  of  P<0.01; F i g .  impossible to assess  activities.  visit  P=0.03),  (G[3]=104.06,  was w i t h o t h e r  closely,  frequency  of  the  attempt  to  pack.  movements of  wolves that  movements  ties  (Fig.  with t h e i r  4 successful dispersals  individually 3).  packs.  In  9 cases,  They  where 2 a n i m a l s  acquired  127  80r  Q UJ UJ z I<  70  <  60  <  Q_ UJ CO  Z <  CO  *  z  <  (5) 44 (8) 58  50  o o  30  (13) 98  20  co »"  (20) 165  (10) 77  40  i= °X UJ 0 X  o  (5) 69  (6) 89  (17) 257 (6) 12  (8) 31  C22) 212 80  10  °  DC  D  J  F  M  2.  December t o M a r c h c h a n g e s  pup, y e a r l i n g , their  packs.  figure portion  in of  J  F  M  D  YEARLING  PUP  Fig.  D  J  F  M  ADULT  sightings  when  and a d u l t w o l v e s w e r e o b s e r v e d a l o n e and s e p a r a t e d  from  Number o f  sightings  in the percentage of  is  given  above t h e c o l u m n s ,  parentheses  d e n o t e s t h e number o f  animals  sightings  during e x t r a - t e r r i t o r i a l  involved.  movements  is  and  the  The  stippled.  128  BOUNDARY OF LA VERENDRYE RESERVE  ?  APPROX. NORTHERN LIMIT OF DEER  *  WOLF PACK TERRITORIES  PACK ORIGIN UNCERTAIN DISPERSAL  EXTRATERRITORIAL EXCURSIONS, SINGLE ANIMALS  Fig. the  3.  The 56 e x t r a - t e r r i t o r i a l  study.  distances.  The e x c u r s i o n s  excursions  are presented  made by s i n g l e w o l v e s  as t h e minimum s t r a i g h t  during line  1 29  a new a s s o c i a t e by an a d j a c e n t lost  all  and formed a pack pack;  wolf  that  after  Dispersals  occurred exclusively  involved  trips,  was  1 case  (March),  killed  Excursion adults  (26),  frequencies  by t h e  frequency  differed  1.0  per  year  prone to  of  per  males  (1  pups  fewer  outside  of  from t h e  and  times. (101, Pups  expected  yearlings  territory  May).  trio-pack  (21),  P<0.01).  s o l o e x c u r s i o n s than  the p a c k ' s  adjacent  expected  radio-tracking  whereas  a  2.1  change o c c u r r e d .  yearlings  G[2]=13.2,  Each  a minimum of  when t h e  (9),  and  4).  i n M a r c h and 1 i n  incomer)  from t h e  respectively),  foray  (Fig.  W o l v e s a c c e p t e d by  significantly  calculated  initiated  initiated  we  and 2  4 males and 3 f e m a l e s  on a v e r a g e .  and 317 months r e s p e c t i v e l y ; adults  journeying;  t h e presumed d o m i n a n t male  (possibly  accepted  d u r i n g which  from m i d - A p r i l t o A u g u s t  p a c k s were b o t h young a d u l t In  extensive  dispersed d e f i n i t i v e l y ,  pre-dispersal  and 2 were  3 probable d i s p e r s a l s  radio-contact  mortalities.  nucleus,  86,  and (  1.1  and  were much more  (3.0  excursions  year). Thirty-eight  winter  period,  classes  (Fig.  territory  forays  but 4).  alone  restricted  of  the  the  seasonal  before mid-February.  primarily  to the w i n t e r seasonal  e x c u r s i o n s d u r i n g the  non-reproductive  patterns  Pups d i d not v e n t u r e  showed no o b v i o u s  15 a d u l t  56 e x c u r s i o n s o c c u r r e d d u r i n g  adults  (24 t o  differed  outside  Yearling  period, pattern.  30 m o n t h s ,  between  the  age  pack's  e x c u r s i o n s were  whereas At  summer p e r i o d  the  least  adult 8 of  involved  from  the  young  known-age  130  PA:  v  TR?  T$X7Vp  iFvfi?  MONTH  Fig.  4.  Chronological  territorial  distributions of the initiation  extra-  e x c u r s i o n s made by s i n g l e p u p s ( P ) , s i n g l e y e a r l i n g s ( Y ) ,  single adults  ( A ) , and packs ( P A ) .  excursions which resulted A measure of r a d i o - t r a c k i n g transmitters)  of  The c l o s e  i n an a c t u a l  symbols denote the  o r assumed d e f i n i t i v e  t i m e (number o f animal-months w i t h  i s presented to facilitate  interpretation.  dispersal. operating  131  animals).  In  summer, I  reproductive adults Also,  I  never  territorial  d i d not  initiate  had e v i d e n c e  movements i n  observe  extra-territorial  that  were m a r k e d ) .  Therefore,  the  implicated single  territory  The d u r a t i o n of for  pups,  13.3 ± 3.2  No d i f f e r e n c e  assumed t h a t  distance  for  of  km f o r  adults.  yearlings  movements  2 1 . 7 ± 2.6  These d a t a  for  indicate  t o be o u t s i d e  caused mainly  P>0.20),  the p a c k ' s  by t h e h i g h e r  e x c u r s i o n s as o p p o s e d t o  adults.  (one-way  and a v e r a g e d  the  the d u r a t i o n or  22.2 ±  tendency  (Fig.  which they  classes  and 2 5 . 5 ± 4.2  higher  territory  r a t e at  for  among age  yearlings,  that  days  t h e minimum s t r a i g h t  e x c u r s i o n s d i d not d i f f e r  pups,  outside  ± (SE)2.6  and 10.3 ± 2.1  Likewise,  (one-way ANOVA, F [ 2 , 5 3 ] = 0 . 3 0 , (SE)5.3  all  1 wolf  wolf.  yearlings,  P>0.20).  extra-  when more than  among age c l a s s e s c o u l d be d e t e c t e d  ANOVA, F [ 2 , 4 0 ] = 0 . 5 4 , line  (i.e.,  e x c u r s i o n s a v e r a g e d 9.1 for  movements.  a pack e n g a g e d i n  summer I  known or presumed  1)  of  was  initiated  l e n g t h of  these  excursions. To examine t h e p o s s i b l e e f f e c t s upon e x t r a - t e r r i t o r i a l table 1).  analyses  Neither  territorial the  tendency  (P<0.01); adults,  movements,  movements of  these  p r e y abundance and sex  conducted contingency  on t h e a g e - s p e c i f i c w i n t e r  p r e y abundance nor  of  I  of  pups  yearlings effects  to  sex  influenced  (P>0.20). leave  were not  observations  Both  interactive  o n l y p r e y abundance was p o t e n t i a l l y  extrafactors  the p a c k ' s  (Table  affected  territory (P>0.10).  effective  In  1 32  (P=0.09).  Among age c l a s s e s ,  significant comparable inclined  (G[2]=45.8, foraying  to t r a v e l  L o c a t i o n s of territorial  territories  that  solitary  outside  cores  the p a c k ' s  However,  (n=359)  frequently  (depicted  in F i g .  the  used t h e  r e p r e s e n t e d the  2),  e d g e s of  the  the  than e x p e c t e d  (Fig.  in  1).  hypothesis  than within  places  in which  cores  (Table  2).  intensively, Altogether,  reveal  a progressive  increase  buffer  zones,  interstices.  and t o  Pack e x t r a - t e r r i t o r i a l  of  use  I  and  observed  within  the  (Table  2).  territories  2-  less  presumably  to a v o i d  pack  Known l o n e w o l v e s showed a d i f f e r e n t cores  being  territory  territories  though these areas  to  expected  and more f r e q u e n t l y  between  extra-  relation  territories.  frequently  interstices even  safest  used t e r r i t o r y  I  3;  zones b o r d e r i n g these  (Table  interactions.  were more  territory  were a n a l y s e d  neighbouring packs,  they  had  wolves would tend t o a v o i d t e r r i t o r y  km wide b u f f e r  they  but y e a r l i n g s  a s s u m i n g homogeneous d i s t r i b u t i o n ) of  highly  pups and a d u l t s  f o r a y i n g pack members l e s s  (i.e.,  was  l o n e w o l v e s and pack members d u r i n g  t o use p r e f e r e n t i a l l y solo  P<0.01);  tendencies,  movements  pack  the d i f f e r e n c e  pattern;  and i n t e r s t i c e s  the a n a l y s i s  less  does  from t e r r i t o r y  not  cores  to  movements  o b s e r v e d 23 i n s t a n c e s (Fig.  of  territorial  movements  5).  exclusively  from 1 December t o  p a c k s engaged i n  extra-  T h e s e e x c u r s i o n s were 30 M a r c h , and many of  observed them  (13)  T a b l e 2. of  Location frequencies  lone wolves,  o f p a c k members d u r i n g s o l o e x t r a - t e r r i t o r i a l  are reported f o r  the t e r r i t o r y  cores,  t h e 2-km w i d e b u f f e r  excursions  zones,  and  and  the  2 interstices  (see methods).  probability  level  A l s o presented are the  {d.f.=2 for  Territory  -tests  of  homogeneity,  and  their  all).  cores  Buffer  zones  Interstices  % area  %fixes  %area  %fixes  %area  %fixes  A  2  P  Pack  members  56  33(57)  24  48(84)  20  19(33)  59.4  <0.01  Lone  Wolves  56  80(148)  24  16(29)  20  4( 8 )  47.0  <0.01  56  57(205)  24  32(113)  20  11(41)  21.6  <0.01  Total  ( ) = sample  size  134  Fig.  5.  study.  The 23 e x t r a - t e r r i t o r i a l Excursions  excursions  made by w o l f  a r e p r e s e n t e d as t h e minimum s t r a i g h t  packs d u r i n g line  the  distances.  1 35  were d e s t i n e d t o d e e r were k i l l e d  wintering  areas.  by w o l v e s d u r i n g t h e s e  were v i c t i m s . from a e r i a l  Given  the  tracking,  exact  As d e p i c t e d by F i g .  distribution  were more i n c l i n e d and t o  Excursion a  low p r e y  winter),  exploit  frequency  abundance  5,  (n=l8,  range=5.5 to  The d u r a t i o n (LP  of  range=2 t o  25),  in  for  packs a s s o c i a t e d  P=0.04).  packs  base a r e a  The minimum  greater  75.7)  than  in  their  i n HP  during (n=5,  straight  (average= 1 2 . 0  U-test[18,5]=72,  P=0.03).  same t r e n d between  was not  with  LP  8 4 ; HP a v e r a g e = 7 . 0  difference  (Mann-Whitney U - t e s t [ 1 3 , 5 ] = 4 7 ,  of  the h i g h prey  range=2 t o  the  be  from  e x c u r s i o n s showed t h e  but  more  carcasses  t o make e x c u r s i o n s  3 0 . 7 ; Mann-Whitney  average=16.5 d a y s ,  deer  deer  e x c u r s i o n s was a l s o  ( a v e r a g e = 2 6 . 1 km, r a n g e = 5 . 0 t o km,  likely  to  60 t r a c k i n g - m o n t h s  2  of  packs c l o s e  was h i g h e r  as compared t o p a c k s  distance  detecting  8 deer  deer.  46 t r a c k i n g - m o n t h s ; X [ 1 ] = 4 . 3 9 , line  of  number c o u l d not  ascertained.  territories  e x c u r s i o n s but  difficulty  the  A minimum of  strongly  areas  days, expressed  P=0.10).  DISCUSSION  Solitary  living,  extra-territorial  To many p e o p l e individual its  from i t s  parent(s)  attempts  " d i s p e r s a l " connotes a natal  to another  to p a i r  movements,  site site  and out at  which  with a c o n s p e c i f i c  of  and  "movement  of  the  it  breeds,  the  dispersal of  home range or a t  opposite  sex  an of least for  1 36  p u r p o s e of to  breeding"  (Bekoff  1 9 7 7 : 7 1 5 ) . The f i r s t  from t h e  present  wolf  be drawn  not  an u n i q u e  study  "one d e c i s i o n - one t r i p "  a p p e a r e d t o be a dynamic and g r a d u a l breaking t i e s  with i t s  pack  over  is  event.  that  in c e n t r a l  that  yearling  forays adult only  per age  wolves  year, (6 out  a fraction  disperse,  initiated  (1983)  of  territorial Age'of influencing Yearlings  frequency  It  and o t h e r  I  of  extra-territorial at  prone  o b s e r v a t i o n s , ^ addressed incentive  for  an  of  which  to  1 s o l o e x c u r s i o n per  the  more  observed  Even among a d u l t s ,  year  the such  extra-  the animals solitary  living  animals likely  was  i d e n t i f i e d as a p r e d o m i n a n t  factor  to  and e x t e r r i t o r i a l  reaching puberty, travel their  solitarily natal  has been s u g g e s t e d t h a t auxiliaries  and t o  territories.  Many  a c t e d as  assistance  food for  wolf  the  spend  with a minimal  the  in providing  movements.  and were c l e a r l y  and young n o n - r e p r o d u c t i v e a d u l t s  a s s o c i a t e d pack members, a p p a r e n t l y pack.  3  age.  forays.  p r o l o n g e d p e r i o d s out of yearlings  one or  study,  were n o n - b r e e d e r s and t h e r e f o r e  what had c o n s t i t u t e d  are  a n i m a l s most  this  on a v e r a g e  7 dispersals).  an a v e r a g e  of  In  of  few  reported  a l t h o u g h most d i s p e r s e d p e r m a n e n t l y  r e c o r d e d . From t h e s e  question  trips.  a  10 months of  Van B a l l e n b e r g h e  extra-territorial  wolf  a p r o t r a c t e d p e r i o d of as  is  Dispersal  d i s p e r s i n g w o l v e s showed a common p a t t e r n  preliminary  was  Alaska,  dispersal  process, with a  months t o a few y e a r s b e g i n n i n g as e a r l y Recently  that  conclusion  of  loosely-  role  in  the  yearlings  pups does  not  1 37  influence et  al.  their  1983; F i g .  yearlings  they  (Harrington  13 i n  Peterson  as  "floater  et  animals",  al.  but  1982;  1984).  they  surviving  and u l t i m a t e l y  parents'  probability  use o t h e r  interest of  these  packmates  to  facilitate  with the  highly-valued  Concomitant w i t h the  effect  peculiarity packs;  status.  survival  and  to  s u b o r d i n a t e wolf or  it  sufficient  of  may e i t h e r  may v o l u n t a r i l y  yearlings  and a d u l t s  age,  wolf  i n HP but  alone  had a g r e a t e r in  the  low  low p r e y  3.2  i n LP  (Messier  evicted  (Zimen  1976,  tendency prey  p a c k s u s e d an a v e r a g e  only  It  is  breeding  abundance movements.  of  to  area  from t h e 1982). live  (Table  and C r e t e  Both  solitarily 1).  6.2 moose p e r  During  100 d a y s  in p r e s s ) .  In  area,  p a c k s were e x p e r i e n c i n g p e r i o d s of  scarcity  which  u n d o u b t e d l y e n c o u r a g e d pack d i s s o c i a t i o n .  circumstances,  dispersal survive  strategy,  forays but  cannot  rather  through a temporary  resource  i n f o r m a t i o n a c q u i r e d d u r i n g these advantageous  subsequently  distribution  i n c l u d i n g the  (e.g.,  immediate failure.  the  food  be a s c r i b e d t o a  as an  a  pack,  low p r e y  these  in  d u r i n g an e x t e n d e d p e r i o d ,  be a c t i v e l y  forage  and t o make e x c u r s i o n s winter,  prey  in  offspring.  i n c r e a s e d pack d i s s o c i a t i o n and e x t r a - t e r r i t o r i a l Without access  typified  a s an a i d  breeder  the  many  c o u l d be  selfishly  in achieving  Harrington  Thus,  keep a m i n i m a l a s s o c i a t i o n w i t h t h e i r  possibly,  the  and Mech  and young n o n - r e p r o d u c t i v e a d u l t s  biologically that  survival  Under  specific  necessity However,  to  the  e x c u r s i o n s may be territory  locations  of  vacancy,  deer  prey  wintering  areas).  138  The g r e a t e r females  to d i s s o c i a t e  territory the  (yearlings  relatively  packs.  tendency  of  yearling  from t h e p a c k , only),  appears  low a v a i l a b i l i t y  Under n u t r i t i o n a l  of  stress,  1976,  and t o  foray  largely  outside  moose c a r c a s s e s more  to  to  the  often  pack members t h a n a r e  1982). These a g g r e s s i v e  the  attributable  females are  s u b o r d i n a t e and h a r a s s e d by o t h e r (Zimen  f e m a l e s and a d u l t  interactions  may  males  result  i n weak pack bonds or even  i n a t e m p o r a r y pack e v i c t i o n .  Not  enough i s  dispersal  that  known a b o u t w o l f  females are  is  However,  not  pre-dispersal  and t o  (Van  their  Ballenberghe  natal  social  receptive  pack or  exploratory habitat travel  unit  great  study appear territorial  wolves to reflects wolf  (and a r e a ) to  return  their  is  high,  vacancy  dispersal  failure  to  as a base  a wolf  territory find  vacant  c o u l d use  from w h i c h t o  can s e t t l e .  many p r e - d i s p e r s a l t r i p s .  t o be i n c o n f l i c t  its  find a  1982).  d i s p e r s i n g w o l v e s may have  before they  area  a new pack d u r i n g  M e s s i e r and B a r r e t t e  with t h i s  This  e x c u r s i o n s were o b s e r v e d i n LP  than  More  its If  to  process  The r e s u l t s view.  as  c a n be  to t h e i r  and/or a s u i t a b l e  form a n u c l e u s of (see  in  be i n t e r p r e t e d  territory  1983). C o n c e i v a b l y ,  distances  may n e c e s s i t a t e  of  alien  excursions  saturation  movements c a n o f t e n  pack  a receptive,  voluntarily  1982).  d u r i n g which  The t e n d e n c y  rejoin  quickly  forays  suggest  a sex-specific difference  e x c l u d e d (Zimen  Extra-territorial  assessed.  to  more i n c l i n e d than m a l e s t o d e p a r t  from the p a c k . strategy  strategy  of  this  extra-  i n HP a r e a  1 39  even  though vacant  press).  However,  areas  the  were more a v a i l a b l e  low p r e y  prompted s u b o r d i n a t e a n i m a l s frequently. vacant  T h i s poor prey  areas  unfavourable  have  likely  confounded the  that  Fritts  and Mech  forays  when t h e y  Minnesota. saturated  abundance to  initiate  rarely  Of  e x c u r s i o n s more t e n d t o make any  interest  observed  studied a newly-protected  The M i n n e s o t a p o p u l a t i o n was, and w o l v e s  there  in  likely  r e p r o d u c t i o n . T h e s e two  results.  (1981)  (Messier  i n LP most  base would a l s o for  i n LP  is  factors  the  fact  pre-dispersal wolf  population  however,  in  not  were e x p e r i e n c i n g an ample  prey  base. The h i g h e r the  b r e e d i n g season  March)  likely  individuals captivity  to  leave  other  1967; Zimen  the  their  end of  the  reproductive  stable,  but  excursions, 1977;  this  March  (Fig.  explanation  low  is  exert  less  d u r i n g the  probable.  signify  were e v e n l y  et  a  to  some Studies  I  have  (Rabb  et  (1984) more  b r e e d i n g s e a s o n . An food s t r e s s an  at  indication  throughout  the  a shortage  of  distributed  in  greater  period al.  15  movements were u n d e r t a k e n  b a s e d on a h i g h e r  which u s u a l l y  4).  Peterson  food a v a i l a b i l i t y  study),  wolves  of  period.  pack members d u r i n g t h i s  extra-territorial  t o and d u r i n g  from 1 F e b r u a r y  pack d u r i n g t h i s  1982). R e c e n t l y ,  winter  packs p r i o r  i n c r e a s i n g tendency  commonly by s u b o r d i n a t e w o l v e s alternative  within  (approxmatively  showed t h a t  reported that  stress  explains  dominance o v e r al.  social  winter. food  the of  Pack  (Mech  from December  a  to  1 40  Solitary  animals  Often,  it  and pack  and Mech  in areas  1981: 4 4 ) .  s u p p o r t e d by q u a n t i t a t i v e knowledge. different  animals,  d i d not  that  surrounding areas.  to  utilize  single  However,  animal  particular  was a v a i l a b l e parallel  the  in the  analysis  wolves used t e r r i t o r y w o l v e s d i d not a v o i d however, packs,  that  but  it  produce a s p a t i a l  apparently  t o my  b a s e d on 25  interstices  noticeably  between animals.  different  east  induced l a r g e l y  observations) 9 beef side  reveals  appeared more by a  of  where b u t c h e r  gut p i l e s ; Fig.  no t r e n d .  4).  cores.  I  If  this  The 2 known  frequently.  this  temporal  offal  see t h e 4 animal  conclude  do not  suggest, with  segregation did  movements  packs undertook e x t e n s i v e to e x p l o i t  deer.  In  winter  one c a s e  movements  (the  is  lone  I  w o l v e s d i d not a v o i d c o n t a c t that  The  from  segregation.  Pack e x t r a - t e r r i t o r i a l Four  (38  (see  4 p r o l o n g e d e x c u r s i o n s to a  territory  appears  available  neighbouring t e r r i t o r i e s  c o r e s more  solitary  lone  evidence  u s e d by s o l i t a r y  t r e n d was  (a minimum of  arrows  excepted,  zone  yet  that  not  of  D i s s o c i a t e d pack members  which undertook  buffer  published  reported here,  was  of  this  locations  u t i l i z e d by p a c k s  not  reveal  interstices  the p e r i p h e r y  frequently.  is  were more f r e q u e n t l y  within  of  less  data  habitat  that  However,  The o b s e r v a t i o n s  territories  distribution  has been p o s t u l a t e d  w o l v e s were p r i m a r i l y Fritts  spat i a l  southermost  that  alien not  141  pack),  the  wolves  be a t r a d i t i o n a l  had been r e p e a t i n g pattern  The low moose d e n s i t y of  c a p t u r i n g deer  in  in  of  movements a t  the  their  each year  study  area,  wintering  w i t h no a p p a r e n t  argued that  malnutrition  neighbouring  initiated  observations  prey.  territorial Such p a t t e r n  instability base  (Kuyt  across  My d a t a  of  suggest  of  wolves  probably  to  trespass  conform to Mech's  a s s o c i a t e d w i t h a low p r e y  habit  ease  pack e x c u r s i o n s were  i n HP.  LP p a c k s were s u b s t a n t i a l l y also  relative  incites  packs  that  wolves  in winter  in  1972; S t e p h e n s o n  LITERATURE  the  These  low p r e y  areas  on a s p a t i a l l y  and James  abundance  longer.  the  (1977)  (1977)  Moreover,  than wolves  s p a c e use r e s e m b l e s relying  with  have  ample  territorial changeable  prey  1982).  CITED  A r m i t a g e KB (1977) S o c i a l v a r i e t y i n t h e y e l l o w - b e l l i e d marmot: a p o p u l a t i o n - b e h a v i o r a l s y s t e m . A n i m . B e h a v . 585-593. B a r a s h DP (1974) The e v o l u t i o n of marmot t h e o r y . S c i e n c e 185: 4 1 5 - 4 2 0 . Bekoff  M (1977)  individual  societies:  Mammalian d i r s p e r s a l and t h e o n t o g e n y behavioral  to  winter.  Mech  more e x c u r s i o n s t h a n p a c k s  e x c u r s i o n s of  weaker  Packs  of  premeditated d e s t i n a t i o n .  territories.  interpretation.  onset  and t h e  areas,  e n c o u r a g e d s u c h pack e x c u r s i o n s . O t h e r shorter  the  what a p p e a r s  phenotypes.  Am. N a t .  111:  a  25:  general of  715-732.  B e k o f f M, D a n i e l s T A , G i t t l e m a n J L (In p r e s s ) L i f e h i s t o r y p a t t e r n s and t h e c o m p a r a t i v e s o c i a l e c o l o g y of c a r n i v o r e s . A n n . Rev. E c o l . S y s t .  a  142  Brown J L (1983) C o o p e r a t i o n - a b i o l o g i s t ' s d i l e m m a . I n : R o s e n b l a t t J S , H i n d e RA, Beer C , B u s n e l M-C ( e d s ) A d v a n c e s i n t h e s t u d y of b e h a v i o r , v o l . 13. Academic P r e s s , New Y o r k . p p . 1-37. Emlen ST (1982) The e v o l u t i o n of h e l p i n g . I. An e c o l o g i c a l c o n s t r a i n t model. II. 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