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UBC Theses and Dissertations

An examination of wildlife policy in Spatsizi Plateau Wilderness Park Pearse, Anthony Dalton 1983

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AN IN  E X A M I N A T I O N S P A T S I Z I  OF  P L A T E A U  W I L D L I F E  P O L I C Y  W I L D E R N E S S  by ANTHONY DALTON PEARSE B . S c , The U n i v e r s i t y of B r i t i s h C o l u m b i a , 1971  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES School o f Community and Regional  Planning  We a c c e p t t h i s t h e s i s as conforming to the r e q u i r e d s t a n d a r d ,  THE UNIVERSITY OF BRITISH COLUMBIA  April, 0  1983  ANTHONY DALTON PEARSE  P A R K  In  presenting  an  advanced  the L i b r a r y I  further  for  of  this  written  thesis  degree shall  agree  scholarly  by h i s  this  at  the U n i v e r s i t y  make  that  it  purposes  for  freely  permission may  representatives. thesis  in p a r t i a l  is  financial  of  British  Date  -t-  Columbia,  British  by  for  gain  Columbia  shall  the  that  not  requirements I  agree  r e f e r e n c e and copying  t h e Head o f  understood  Jbap<H" tnierrf'tff "~  2075 Wesbrook P l a c e V a n c o u v e r , Canada V6T 1W5  of  for extensive  permission.  The U n i v e r s i t y  of  available  be g r a n t e d  It  fulfilment  of  this  be a l l o w e d  or  that  study. thesis  my D e p a r t m e n t  copying  for  or  publication  without  my  II  A B S T R A C T  My o b j e c t i v e i n t h i s t h e s i s  i s to compare the c o r r e l a t i o n  between p o l i c y i n t e n t and p o l i c y outcome w i t h r e s p e c t to w i l d l i f e management  in Spatsizi  Council e s t a b l i s h i n g  Plateau Wilderness  The O r d e r - i n -  the park a r t i c u l a t e s a management  which i s t o a l l o w " n a t u r a l communities progressions  Park.  policy  to remain i n t a c t and the  o f n a t u r a l systems to proceed w i t h o u t a l t e r a t i o n . "  The study begins  by i n t e r p r e t i n g the p o l i c y statement to mean t h a t  no human a c t i o n w i l l  be p e r m i t t e d w i t h i n t h e park t h a t w i l l  the e c o l o g i c a l o r d e r of e x i s t i n g w i l d l i f e communities. quent a n a l y s i s  is divided into four stages.  disrupt  The subse-  F i r s t , I i d e n t i f y the  key, o p e r a t i n g p r i n c i p l e s t h a t govern n o r t h e r n b o r e a l f o r e s t such as t h a t c h a r a c t e r i z i n g S p a t s i z i . guidelines  Second,  I formulate  from these i d e n t i f i e d e c o l o g i c a l p r i n c i p l e s .  lowed by a r e v i e w of p r e v a i l i n g management  ecosystems  management  This  is  p r a c t i c e s i n t h e park.  I a s s e s s the p r e v a i l i n g p r a c t i c e s i n terms of t h e i r c o n s i s t e n c y my d e r i v e d g u i d e l i n e s  folLast  with  i n o r d e r t o d e t e r m i n e t o what e x t e n t p o l i c y ob-  j e c t i v e s are being met. The r e s u l t s of t h i s a n a l y s i s guidelines  revealed that of s i x  management  developed f o r S p a t s i z i w i l d l i f e p o l i c y , o n l y one was  l y being implemented by park managers. w i l d l i f e management  in Spatsizi  The c o n c l u s i o n i s  present-  that current  Park i s not being c a r r i e d out on an  e c o l o g i c a l l y sound b a s i s and, t h e r e f o r e , i s not a c h i e v i n g , and i s to a c h i e v e , the o b j e c t i v e s of the s t a t e d p o l i c y .  E m p i r i c a l support  l e n t t o t h i s c o n c l u s i o n by the apparent d e c l i n e o f s e v e r a l l a t i o n s w i t h i n the p a r k .  unlikely  ungulate  is popu-  My recommendations  i n c l u d e a temporary s u s p e n s i o n o f a l l  b u n t i n g w i t h i n and a d j a c e n t to the park u n t i l studies  the n e c e s s a r y  have been conducted and the e s s e n t i a l c r i t e r i a f o r  biological enabling  the h a r v e s t to occur i n accordance w i t h the p o l i c y mandate have been met.  An e s s e n t i a l  element here i s t h a t a w i l d l i f e r e f u g e  v i a b l e p o p u l a t i o n s of a l l b i g game s p e c i e s  i n the S p a t s i z i  containing r e g i o n be  e s t a b l i s h e d w h e r e i n no development or e x p l o i t a t i o n o f w i l d l i f e be allowed to take p l a c e .  iv  TABLE OF CONTENTS page Abstract  i i  L i s t of T a b l e s  vii  L i s t of  vi  Figures  Acknow!edgements  vi i i  Bib"! i o g r a p h y  105  CHAPTER I  Introduction  1.1  Study Scope and O b j e c t i v e s  1  1.2  Methodology  8  CHAPTER II  E c o l o g i c a l "Rules of Order"  2.1  Introduction  2.2  The O r g a n i z a t i o n of Ecosystems I n t e r c o n n e c t e d n e s s and Order  2.3  N o n - l i n e a r i t i e s and M u l t i p l e E q u i l i b r i a  13  2.4  Dynamic V a r i a b i l i t y  16  2.5  Uncertainty  17  CHAPTER III  10 -  12  P r e d a t o r - P r e y Systems  3.1  Introduction  19  3.2  P r e d a t o r - P r e y Dynamics  21  3.3  P r e d a t o r Response and A l t e r n a t e Prey  25  CHAPTER IV  The E f f e c t s of Hunting  4.1  Introduction  4.2  Q u a n t i t a t i v e Aspects of Maximizing  4.3  Q u a l i t a t i v e Impacts o f the H a r v e s t  CHAPTER V 5.1  34 Harvests  35 45  S p a t s i z i - A D e s c r i p t i o n of the W i l d l i f e Component  S p a t s i z i W i l d l i f e Communities - An Overview  50  V  CHAPTER VI 6.1  Ecological Guidelines f o r Spatsizi W i l d ! i f e Management  G u i d e l i n e s f o r W i l d l i f e Management  CHAPTER V I I  C u r r e n t W i l d l i f e Management P r a c t i c e s in S p a t s i z i Park - D e s c r i p t i o n and A n a l y s i s  7. 1  Introduction  7. 2  Calculation-of the Harvestable  7. 3  7..4  7..5  7..6  7..7  61  70 Surplus  72  7.2.1  The Rule  72  7.2.2  Present Regulation of the Harvest  72  7.2.3  Deficiencies in Current Practices of Establishing Harvest Levels  77  Age/Sex D i s t r i b u t i o n o f t h e P o p u l a t i o n  81  7.3.1  The Rule  81  7.3.2  Present Regulation of the Harvest  81  7.3.3  Consistency with the Guideline  81  Management U n i t s  81  7.4.1  The Rule  81  7.4.2  P r e s e n t Management A p p r o a c h  81  7.4.3  Consistency with the Guideline  86  Pulse Harvesting  88  7.5.1  The Rule  88  7.5.2  Present Practice  89  7.5.3  Consistency with the Guideline  89  Predator  Management  91  7.6.1  The Rule  91  7.6.2  Present Practice  91  7.6.3  Consistency with the Guideline  93  The E s t a b l i s h m e n t o f Benchmark P o p u l a t i o n s  94  7.7.1  The Rule  94  7.7.2  Present Practice  94  7.7.3  Consistency with the Guideline  94  CHAPTER V I I I  C o n c l u s i o n s and Recommendations  8,.1  Synthesis  8,.2.  Recommendations  ;  97  LIST OF FIGURES Page 1.1  L o c a t i o n Map of S p a t s i z i P l a t e a u W i l d e r n e s s Park  2  1.2  R e g i o n a l S e t t i n g of S p a t s i z i W i l d e r n e s s Park  3  2.1  D e n s i t y dependent p o p u l a t i o n dynamics  14  2.2  Simple r e c r u i t m e n t curve  14  3.1  D e n s i t y dependent p o p u l a t i o n dynamics  23  3.2  Prey s p e c i e s r e c r u i t m e n t curve  23  3.3  Time l a g s i n p r e d a t o r - p r e y systems  26  3.4  E f f e c t s of e n v i r o n m e n t a l v a r i a b i l i t y on r e c r u i t m e n t  27  4.1  Prey r e c r u i t m e n t curve  36  4.2  E f f e c t s o f the h a r v e s t on r e c r u i t m e n t f o r subsequent y e a r s  38  4.3  Annual and seasonal t r e n d s i n p o p u l a t i o n dynamics e x c l u d i n g random e n v i r o n m e n t a l e f f e c t s  39  4.4  Annual c y c l e o f u n g u l a t e p o p u l a t i o n dynamics showing q u a l i t a t i v e e f f e c t s of h a r v e s t i n g  39  4.5  E f f e c t s of random e n v i r o n m e n t a l events on annual u n g u l a t e p o p u l a t i o n c y c l e s i n n a t u r a l and hunted  42  Plateau  states 4.6  C a r i b o u p o p u l a t i o n d e c l i n e i n B r i t i s h Columbia  44  5.1 5.2  N o r t h e r n b o r e a l f o r e s t ecosystem f o o d c h a i n P o p u l a t i o n and h a b i t a t v a r i a b l e s a f f e c t i n g d e n s i t y o f moose  52 53  5.3  D e n s i t y dependent r e c r u i t m e n t curve f o r c a r i b o u  58  7.1  S p a t s i z i Wilderness  71  7.2  P o p u l a t i o n t r e n d s of c a r i b o u i n S p a t s i z i  7.3  W i l d l i f e Management U n i t s s u r r o u n d i n g  7.4  S p a t s i z i Park showing o v e r l a p p i n g G u i d e / o u t f i t t e r t e r r i t o r i es  85  7.5  Management U n i t boundaries superimposed on g u i d e / o u t f i t t e r t e r r i t o r i e s and S p a t s i z i Park  85  Park - P h y s i c a l F e a t u r e s Park  Spatsizi  78 Park  84  YLl  LIST OF TABLES  7.1  Large mammal p o p u l a t i o n s i n S p a t s i z i  Park  7.2  H a r v e s t o f b i g game from S p a t s i z i  7.3  P r o v i n c i a l h a r v e s t o b j e c t i v e s f o r b i g game s p e c i e s  90  8.1  C o n f o r m i t y of S p a t s i z i management p r a c t i c e s w i t h d e r i v e d management g u i d e l i n e s  99  Park 1975-1980  75 82  A C K N O W L E D G E M E N T S  I owe my deepest sense o f g r a t i t u d e to the u n f a i l i n g c a p a c i t y of I r v i n g  Fox to always keep t h i s study on t r a c k by t u r n i n g on the  l i g h t a t the end o f the t u n n e l d u r i n g p e r i o d s o f d a r k n e s s .  His b e l i e f  i n the worth of the p r o j e c t and t h a t i t would someday be c o m p l e t e d , a f a i t h sometimes g r e a t e r than my own, p r o v i d e d much o f the  necessary  s t r e n g t h and guidance to a c h i e v e the t a s k .  I owe much a l s o t o the  rigourous,  Rees whose uncanny  seal p e l - w i e l d i n g t a l e n t s o f B i l l  ability,  to s e p a r a t e the wheat from the c h a f f c o n t r i b u t e d g r e a t l y to a more meaningful f i n a l  product.  A d d i t i o n a l l y , the study c o u l d not have been  completed w i t h o u t the unwavering p a t i e n c e o f the woman o f my L i n d a A l e x a n d e r , and my good f r i e n d and mentor, B i l l  HorswilT.  life, Finally,  and by way o f d e d i c a t i o n , I cannot submit t h i s t h e s i s w i t h o u t m e n t i o n i n g t h r e e small s o u l s who have always been i n the back o f my mind d u r i n g this exercise:  E z r a , E r i n , and Jeremy, f o r whom t h i s i s w r i t t e n i n the  hopes t h a t the r i c h n e s s o f w i l d l i f e r e s o u r c e s will old.  i n B r i t i s h Columbia today  remain u n d i m i n i s h e d l o n g a f t e r t h e y , and t h e i r c h i l d r e n , have grown  Deep  in  the  pervasive that is  human need  makes  unconsciousness  for  sense.  always  one  a But  step  is  logical the  beyond  a  universe real  universe  logic.  Frank H e r b e r t  Perhaps of  all  the our  most  vulnerable  and  resources.  Durward A l l e n  fragile  C H A P T E R  I  Introduction  1.1  Study Scope and O b j e c t i v e s The purpose o f t h i s t h e s i s  i s to examine the c o n s i s t e n c y  between  s t a t e d p o l i c y and p r e v a i l i n g w i l d l i f e management p r a c t i c e s i n S p a t s i z i  Wilder-  ness P a r k , an area o f some 3,000 square m i l e s i n n o r t h e r n B r i t i s h C o l u m b i a . U n l i k e most B.C.  parks, t h i s area  was bestowed w i t h p a r t i c u l a r i n s t r u c t i o n s  f o r the management o f i t s w i l d l i f e r e s o u r c e s when, i n 1976, i t was t r a n s f e r r e d from undesignated Crown l a n d .  Indeed, i t was the r e c o g n i t i o n o f the compara-  t i v e l y r i c h and d i v e r s e endowment o f w i l d animal p o p u l a t i o n s t e r r i t o r y t h a t a p p a r e n t l y l e d t o the c r e a t i o n of the park  i n the S p a t s i z i  itself.  The area i n and around the S p a t s i z i P l a t e a u i s one o f the few, i f not the o n l y , s i g n i f i c a n t l y l a r g e t r a c t s o f r o l l i n g s u b - a l p i n e upland and a l p i n e tundra i n the p r o v i n c e . i m p o r t a n t b i g game s p e c i e s t o r i c a l l y , abundant.  P o p u l a t i o n s o f almost a l l B r i t i s h  Columbia's  p l u s many non-game s p e c i e s a r e , or have been h i s -  Most prominent o f t h e s e p o p u l a t i o n s , i s perhaps  the  Figure  1.1  L o c a t i o n Map o f  Spatsizi  Plateau Wilderness  Park  ure 1 . 2  Regional  S e t t i n g of S p a t s i z i  Plateau Wilderness  Park  4  woodland c a r i b o u f o r whom the S p a t s i z i  serves  range, r u t t i n g a r e a , and m i g r a t i o n c o r r i d o r .  as a c a l v i n g ground,  summer  The a r e a a p p a r e n t l y serves  as  home f o r a c a r i b o u p o p u l a t i o n e s t i m a t e d t o be between 2,000 and 3,000 a n i m a l s . The park was c r e a t e d as a C l a s s A p r o v i n c i a l park by O r d e r - i n C o u n c i l No. 3756.  The i n c l u s i o n o f s p e c i a l  instructions  i n the O r d e r - i n -  C o u n c i l s e t the o p e r a t i o n a l o b j e c t i v e s f o r w i l d l i f e management w i t h i n the park d i s t i n c t i v e l y a p a r t from t h o s e p r e v a i l i n g throughout the p r o v i n c e , i n c l u d i n g o t h e r p a r k s . makes  the remainder c f  The preamble to the  Order-in-Counci1  i t c l e a r t h a t the park has w i l d l i f e v a l u e s of s p e c i a l s i g n i f i c a n c e  B r i t i s h Columbians  to  and t h a t e x t r a o r d i n a r y a t t e n t i o n to p r e s e r v a t i o n and manage-  ment must o c c u r : " W i t h i n the b o u n d a r i e s o f B r i t i s h Columbia are encompassed some r e g i o n s i n which the p o p u l a t i o n s i z e and d i v e r s i t y of w i l d l i f e s p e c i e s i s of g r e a t b i o l o g i c a l s i g n i f i c a n c e and o f i n s p i r a t i o n a l and e d u c a t i o n a l v a l u e to the p u b l i c ; . . . S u c h unique w i l d l i f e areas r e q u i r e e x c e p t i o n a l p r o t e c t i o n and management to ensure t h a t the v a l u e s a s s o c i a t e d w i t h the w i l d l i f e are r e t a i n e d and not p e r m i t t e d to degenerate i n q u a l i t y , " The uniqueness  o f t h i s preamble l i e s i n i t s emphasis  upon the b i o -  l o g i c a l , i n s p i r a t i o n a l , and e d u c a t i o n a l v a l u e o f w i l d l i f e f o r people and  its  complete o m i s s i o n  in-  o f any r e f e r e n c e t o consumptive o r u t i l i t a r i a n v a l u e s ;  deed, i t i s unique i n p r o v i n c i a l w i l d l i f e p o l i c y s t a t e m e n t s . the maintenance o f the q u a l i t y o f t h e s e v a l u e s a d m i n i s t r a t i o n o f the park.  is  Furthermore,  to be a p o l i c y goal  f o r the  P o l i c y makers a t the time saw c l e a r l y t h a t  ' e x c e p t i o n a l ' measures would be r e q u i r e d t o a c h i e v e t h i s  goal.  The management o b j e c t i v e s f o r the park were l a i d out even more e x pressly  i n the  Order-in-Counci1: f  "The a r e a w i l l be m a i n t a i n e d as a w i l d e r n e s s landscape i n which n a t u r a l communities are p r e s e r v e d i n t a c t and the p r o g r e s s i o n s o f the n a t u r a l s y s t e m s may p r o c e e d w i t h o u t a l t e r ation. H u n t i n g and f i s h i n g w i t h i n s u s t a i n e d yield limits is permissible " In t h i s statement  l i e s the f i r s t h i n t of a possible inconsistency  with the preamble - the p r o v i s i o n f o r consumptive Nonetheless, and  the mandate f o r managers i s r e l a t i v e l y c l e a r :  f i s h i n g may  communities  be p e r m i t t e d  are preserved  remains  unperturbed.  concern  lies.  of view,  use of w i l d l i f e while  i t must occur w i t h i n l i m i t s such  i n t a c t and  I t i s in this p o l i c y statement  E x a c t l y what, from both  the unaltered progression of natural  i n t a c t and  perhaps  i n the present debate  and  B.C.  Some o f t h o s e Parks  contravening  t h a t some b i g game p o p u l a t i o n s a r e , o r a p p e a r s h o u l d be s u s p e n d e d  until  a dominant  Branch,  argue  alteration.  are not adequately  that  low  the management o b j e c t i v e s Others  argue that  enough r e s e a r c h has been done to  level  s i s t e n t with the mandate r e q u i r i n g t h a t n a t u r a l systems without  U n l i k e the second  use  involved, including  t o be, d e c l i n i n g and  t h a t , r e g a r d l e s s o f p o p u l a t i o n l e v e l s , any  groups,  and i n d i v i d u a l s  harvest l e v e l s that are c o n s i s t e n t with the Order-in-Counci1. argue  con-  research  over  t h e r e f o r e , are c o n s i s t e n t with the O r d e r - i n - C o u n c i 1 .  hunting  allow  Several o r g a n i z a t i o n s i n c l u d i n g the  o f the park as a t r o p h y h u n t i n g r e s e r v e .  and,  to  f u e l e d the c u r r e n t  c o n s e r v a t i o n i s t s , and a v a r i e t y o f p r i v a t e i n t e r e s t g r o u p s  l e v e l s of hunting can occur without  my  the p u b l i c ' s point  p e r t i n e n t government agencies, p r i v a t e f o u n d a t i o n s , academic  F i s h and W i l d l i f e Branch  systems  systems?  t r o v e r s y over S p a t s i z i Park management.  t h e B.C.  that natural  that the focus of  t h e m a n a g e r ' s and  I t i s t h i s key q u e s t i o n t h a t has  to varying degrees  hunting  that the progression of natural  d o e s i t mean t o p r e s e r v e n a t u r a l c o m m u n i t i e s  are embroiled  resources.  p o s i t i o n which  determine  Still  of hunting be a l l o w e d  others  is inconto  proceed  i m p l i e s t h a t managers  p r o t e c t i n g population stocks, this t h i r d p o s i t i o n holds  6  that there itself.  i s an  i n c o n s i s t e n c y or c o n t r a d i c t i o n i n the  I t i s c o n t e n d e d b y t h o s e who  sustained y i e l d or not, communities. and  i s simply  Order-in-Counci1  hold t h i s position that hunting,  incompatible  with  Some a l t e r a t i o n , h o w e v e r s e e m i n g l y  p o s s i b l y p r e c i p i t a t e major, unintended  the i n t e g r i t y of  trivial, will  change.  ment p r a c t i c e s and  on w h a t i s m e a n t by " p r e s e r v i n g  "allowing  natural  systems to proceed  t o have done so i n t h e O r d e r - i n - C o u n c i 1 implementation Spatsizi  Wilderness  this policy statement. public controversy  objectives  to reach natural  without  s o m e common communities  alteration".  The  1980  Failure proper  Parks Branch document e n t i t l e d  Park Master Plan s i m i l a r l y f a i l e d to elaborate  The  i t has  the s t a t e d  i s a major d e f i c i e n c y f o r the  of park w i l d l i f e p o l i c y .  Plateau  inevitably result  the congruency between p r e v a i l i n g manage-  the p o l i c y mandate i t i s necessary  d e f i n i t i o n or accord i n t a c t " and  to assess  natural  In o t h e r w o r d s , m a n a g e r s ,  because o f t h i s c o n t r a d i c t i o n , cannot p o s s i b l y hope to a c h i e v e In a t t e m p t i n g  whether  i m p o r t a n c e o f t h i s l i e s n o t so much i n t h e  stimulated  as  on  great  i t does i n the very p o s s i b l e , r e a l  ( i . e . b i o l o g i c a l ) c o n t r a d i c t i o n that might e x i s t i n the mandate which would not allow managers to achieve The  Master Plan  policy objectives.  i n t e r p r e t s t h e l e g i s l a t i o n as  follows:  "The i n t e n t o f t h i s O r d e r - i n - C o u n c i 1 is clear r e c r e a t i o n a l u s e wi11 b e p r o v i d e d i n t h e p a r k . . . I t i s t h e r e f o r e t h e g o a l o f t h e P a r k s and Outdoor Recreation D i v i s i o n within Spatsizi Plateau Wilderness Park to conserve wilderness lands c a p e , and n a t u r a l p r o c e s s e s w h i l e p r o v i d i n g f o r q u a l i t y , d i s p e r s e d , low i m p a c t , r e c r e a t i o n opportunities within those landscapes." (Master I have provided  P l a n , p a g e 6,  emphasis i n the above q u o t a t i o n  d i s t i n c t i o n between the p o l i c y s t a t e m e n t and The  emphasis  b e c a u s e an  fying of administrative  important  t h e M a s t e r P l a n m u s t be made.  O r d e r - i n - C o u n c i 1 d o e s n o t - i s a y t h a t r e c r e a t i o n a l u s e wi 11  only that i t is permissible.  added)  be  provided,  T h i s i n t e r p r e t a t i o n m u s t be v i e w e d as a c o d i -  i n t e n t to ensure continued  r e c r e a t i o n a l use of  the  park.  I do n o t , however, propose  t o d e a l f u r t h e r w i t h the Master P l a n i n  t h i s e x a m i n a t i o n a s , a p a r t from the d i s t i n c t i o n noted above, i t adds f u n d a m e n t a l l y s i g n i f i c a n t t o the O r d e r - i n - C o u n c i 1 .  This  nothing  l a t t e r remains  the  c h i e f f o c u s o f my a n a l y s i s . To begin w i t h , I i n t e r p r e t the g e n e r a l Council  to be e s s e n t i a l l y " p r e s e r v a t i o n i s t "  i n t e n t of the O r d e r - i n -  in character.  By t h i s  t h a t the s p e c i a l c h a r a c t e r i s t i c s o f S p a t s i z i w i l d l i f e r e s o u r c e s c l e a r l y r e c o g n i z e d by p o l i c y makers as worthy of p r e s e r v i n g natural be.  s t a t e , w i t h e x t r a o r d i n a r y p r o t e c t i v e measures  I mean  have been  them i n t h e i r  to be a p p l i e d i f need  T h e i r c o n t i n u e d e x i s t e n c e i s t o have h i g h e r p r i o r i t y than t h e i r e x p l o i t a -  t i o n by man. The term " n a t u r a l c o m m u n i t i e s "  I take to mean a l l p o p u l a t i o n s  w i l d animals and v e g e t a t i o n t h a t e x i s t as an i n t e r a c t i n g a s s o c i a t i o n t a i n geographical I  l o c a t i o n , i n t h i s case the S p a t s i z i  area.  of  in a cer-  The word " i n t a c t "  i n t e r p r e t to mean t h a t a l l c h a r a c t e r i s t i c s o f any b r e e d i n g p o p u l a t i o n must  be p r e s e r v e d so t h a t the h e a l t h and r e p r o d u c t i v e p o t e n t i a l of each  remains  u n d i m i n i s h e d and the p r e s e n t community s t r u c t u r e i n terms of the b a l a n c e between w i l d l i f e p o p u l a t i o n s  remains  e s s e n t i a l l y the same.  " A l l o w i n g the progression alteration"  of n a t u r a l systems t o proceed w i t h o u t  I i n t e r p r e t s i m p l y as a d i r e c t i v e t h a t t h e r e s h a l l  f e r e n c e i n whatever n a t u r a l f o r c e s f o r change a f f e c t S p a t s i z i 'progressions  of natural systems'  wildlife.  would i n c l u d e s t o c h a s t i c e n v i r o n m e n t a l  such as f o r e s t f i r e s , f l o o d s , e t c . ; s e q u e n t i a l cession  be no i n t e r -  change such as e c o l o g i c a l  i n both v e g e t a t i o n and w i l d l i f e ; or a l t e r a t i o n s  i n community  Such events suc-  structure  t h a t might be caused through p r e d a t i o n , c o m p e t i t i v e e x c l u s i o n , and o t h e r i n t e r s p e c i f i c and i n t r a s p e c i f i c phenomena. With the above i n t e r p r e t a t i o n s p l a c e d on the wording o f the O r d e r in-Counci1  is  i t possible  to conduct h u n t i n g and f i s h i n g a c t i v i t i e s w i t h i n  o confines  of the park such that the p r e s e r v a t i o n i s t mandate i s not  C e r t a i n l y , i t i s p o s s i b l e t o v i s u a l i z e t h a t one  moose, one  r a i n b o w t r o u t m i g h t be t a k e n  a f f e c t i n g the  without  structure or a l t e r i n g natural some l e v e l o f h a r v e s t to occur.  How,  i n a n y way  systems.  On  the other  t h e n , m i g h t one  are t e c h n i c a l l y consistent with  o f c o m m u n i t y i n t e g r i t y and  the progression  If i t i s possible to achieve as a l l o w  some c o n s u m p t i v e use,  lowed or i n f o r m a t i o n to s a f e l y f u l f i l l  My  are there  the  and  i s i t being  applied  fishing,  preservation  w i t h i n the l i m i t s set,  then  guidelines  t o be  fol-  employ And,  if  i n the c u r r e n t management  analysis is directed at attempting  of consistency  begin  systems?  f o r i n t h e i r r e s o l u t i o n l i e s the attainment  t i v e ; the determination  to resolve  o f my  b e t w e e n p o l i c y and  of  these  principal  objec-  practice.  Methodology As  will  system w i l l  t h e o b l i g a t i o n s s e t o u t by t h e O r d e r - i n - C o u n c i 1 ?  Spatsizi wildlife?  1.2  preservation  is clearly  t o be m u s t e r e d t h a t p a r k w i l d l i f e m a n a g e r s c a n  such e x i s t s , to what extent  questions,  of natural  one  community  determine at what l e v e l hunting  even i f l e g a l l y permitted,  as w e l l  g r i z z l y , or  hand t h e r e  at which a l t e r a t i o n s to the natural  violated?  I have taken  the Order-in-Counci1  to mean t h a t n a t u r a l  remain the dominant r e g u l a t o r y mechanisms a f f e c t i n g S p a t s i z i  it follows these  t h a t any management p o l i c y a d o p t e d must have m i n i m a l  processes  and,  an u n d e r s t a n d i n g  Accordingly, general  parts:  wilderness  an  wildlife,  impact upon  f u r t h e r , must adhere to the fundamental p r i n c i p l e s which  g o v e r n t h e s y s t e m ' s s t a b i l i t y and incorporate  processes  my  o f how  persistence. natural  P o l i c i e s so f o r m u l a t e d  s y s t e m s p e r s i s t and f l o u r i s h .  a n a l y s i s of the problem can  i d e n t i f i c a t i o n of the  important  ecosystems of the kind found i n northern  t h e s i s of the p r i n c i p l e s i n t o management g u i d e l i n e s :  must  be d i v i d e d i n t o  operative  four  principles in  B r i t i s h Columbia; a that accord  with  the  syn-  1 d i r e c t i v e of the O r d e r - i n - C o u n c i 1 ;  a review of p r e s e n t p r a c t i c e s and p o l i c i e s  a p p l i e d by park a d m i n i s t r a t i o n ; and, f i n a l l y , a c o m p a r a t i v e a n a l y s i s p r e v a i l i n g s i t u a t i o n u s i n g the d e r i v e d g u i d e l i n e s congruency  as c r i t e r i a f o r  between p o l i c y i n t e n t and p o l i c y outcome ( i . e . p r e s e n t  of  the  determining practice).  10  C H A P T E R II  E c o l o g i c a l "Rules o f O r d e r "  2.1  Introduction I f n a t u r a l p r o c e s s e s a r e t o r e g u l a t e e v e n t s w i t h i n t h e park boun-  d a r i e s i t f o l l o w s t h a t w h a t e v e r management a c t i v i t i e s a r e undertaken  should  not v i o l a t e t h e fundamental p r i n c i p l e s t h a t g o v e r n t h e s t a b i l i t y and s u c c e s sion o f ecosystems.  In o r d e r t o make t h i s d e t e r m i n a t i o n  i tis first  neces-  s a r y t o i d e n t i f y and d e s c r i b e t h o s e p r i n c i p l e s o f w i l d e r n e s s e c o s y s t e m s t h a t influence the composition,  s t r u c t u r e , and dynamics o f w i l d l i f e c o m m u n i t i e s .  This chapter serves the function o f e l u c i d a t i n g these relevant p r i n c i p l e s . F i r s t , i t s h o u l d be noted t h a t w h i l e t h e term ' w i l d l i f e ' p r o p e r l y i n c l u d e s a l l forms o f l i f e , p l a n t and a n i m a l , t h a t r e m a i n  undomesticated,  u n c u l t i v a t e d , o r o t h e r w i s e e s s e n t i a l l y ' u n a l t e r e d by man, my f o c u s i s on t h e dominant w i l d l i f e s p e c i e s r e p r e s e n t a t i v e o f n o r t h e r n B r i t i s h C o l u m b i a - t h e so c a l l e d ' b i g game' s p e c i e s .  These i n c l u d e g r i z z l y b e a r , b l a c k b e a r , moose,  m o u n t a i n g o a t , S t o n e ' s sheep, and woodland c a r i b o u . d e f i n e as t r a c t s o f n a t u r a l l a n d s c a p e  The term ' w i l d e r n e s s ' I  i n t h e o r d e r o f hundreds t o thousands  o f s q u a r e m i l e s i n a r e a l e x t e n t t h a t remain g e n e r a l l y u n d i s t u r b e d o r unmarked by human i n c u r s i o n . A ' w i l d e r n e s s ecosystem'  i s the c o m b i n a t i o n  of  t h i s n a t u r a l environment w i t h i t s a p p u r t e n a n t w i l d l i f e endowment - the d w e l l e r s and the d w e l l i n g , i n o t h e r words. Our knowledge o f w i l d e r n e s s ecosystems  is in i t s infancy. Likely,  i t w i l l be decades b e f o r e e c o l o g i s t s have answered some of the  fundamental  q u e s t i o n s c o n c e r n i n g the dynamic p r o c e s s e s and s t a b i l i t y c h a r a c t e r i s t i c s of these natural environments. o f a l l ecosystems  One t h i n g t h a t i s c l e a r now i s t h a t the b e h a v i o u r  i s sometimes d r a m a t i c a l l y u n p r e d i c t a b l e .  Much o f t h i s  u n c e r t a i n t y w i l l never be reduced because of the c o m p l e x i t y and v a r i a b i l i t y inherent in nature.  Resource managers, as a consequence,  to o p e r a t e w i t h complete i n f o r m a t i o n .  can never e x p e c t  What w i l l l i k e l y be more v a l u a b l e i s  the c a p a b i l i t y to manage e f f e c t i v e l y i n the f a c e o f i n c o m p l e t e i n f o r m a t i o n , the a b i l i t y t o s e l e c t what i n f o r m a t i o n i s c r i t i c a l and what i s i r r e l e v a n t , and the a b i l i t y t o d e s i g n and implement r e s i l i e n t management p o l i c i e s so t h a t the c o s t s o f unexpected  change a r e m i n i m i z e d .  T h e r e i s , however, an emerging body o f t h e o r e t i c a l and e m p i r i c a l e c o l o g i c a l c o n c e p t s t h a t have a p p a r e n t r e l e v a n c e f o r the 'management' o f n a t u r a l systems.  These c o n c e p t s o r p r i n c i p l e s seem to be a b l e to produce  q u a l i t a t i v e l y a c c u r a t e ' p e r c e p t u a l windows' on how n a t u r a l b i o l o g i c a l s y s tems s t a b i l i z e and adapt to t h e many f o r c e s o f change.  T h e s e 'windows', o r  m o d e l s , can o f t e n i l l u m i n a t e the key v a r i a b l e s t h a t most s i g n i f i c a n t l y a f f e c t the system and, t h u s , p r o v i d e some c l u e s as to what s t r u c t u r a l ' l e v e r s ' may be a v a i l a b l e t o t h e r e s o u r c e manager f o r e x p l o i t a t i v e , p r o t e c t i v e , o r r e s t o r a t i v e management s t r a t e g i e s . I t i s t h e s e p r i n c i p l e s t h a t c o m p r i s e t h i s window on w i l d e r n e s s ecosystems  to which I t u r n n e x t .  2.2  The O r g a n i z a t i o n of Ecosystems - I n t e r c o n n e c t e d n e s s  and O r d e r  Recent advances i n e c o l o g y have made i t a b u n d a n t l y systems are not comprised  c l e a r t h a t eco-  o f random and u n r e l a t e d p a r t s b u t , r a t h e r , s p e c i f i c  s e t s o f d i f f e r e n t i a t e d and h i g h l y i n t e g r a t e d p a r t s .  I t i s not j u s t the quan-  t i t a t i v e a s p e c t , such as the number o f s p e c i e s or the s p e c i e s d i v e r s i t y i n a p a r t i c u l a r community which i s the key to e c o s y s t e m s t a b i l i t y but the q u a ! i t a t i v e n a t u r e o f how the p a r t s f i t t o g e t h e r ( H o l l i n g and G o l d b e r g , s p i t e o f the commonly c i t e d d i c t u m " e v e r y t h i n g i s c o n n e c t e d  1971).  In  to e v e r y t h i n g  e l s e " , s t u d i e s o f ecosystems s u g g e s t t h a t the c o n n e c t i o n s between the e l e ments t h a t p r o d u c e the d i s t i n c t i v e o r g a n i z a t i o n o f the system a r e l i m i t e d . In o t h e r words, "ecosystems e x h i b i t p a t t e r n s o f c o n n e c t i o n s assemblies that are t i g h t l y connected w i t h i n themselves,  r e s u l t i n g i n sub-  but l o o s e l y con-  nected to o t h e r s " ( H o l l i n g e t a l , 1978, p.27; see a l s o Simon, 1973).  From  the p e r s p e c t i v e o f management, t h e r e f o r e , i t becomes i m p o r t a n t to d e v e l o p  the  c a p a b i l i t y not o f m e a s u r i n g a l l v a r i a b l e s b u t , r a t h e r , i d e n t i f y i n g the key l i n k s or connections  i n the system t h a t d e f i n e i t s dominant p r o p e r t i e s .  T h i s d e g r e e of component i n t e g r a t i o n o r ' i n t e r c o n n e c t e d n e s s the system c o m p r i s e s  what G o l d s m i t h  in  terms "an o r d e r w h i c h i s measured i n terms  o f the i n f l u e n c e o f the whole o v e r the sum o f the p a r t s " ( G o l d s m i t h , As used by G o l d s m i t h ,  1  1978).  the c o n c e p t o f ' o r d e r ' p e r t a i n s to the q u a l i t a t i v e f e a -  t u r e o f e c o l o g i c a l systems t h a t combines the n o t i o n s o f d i v e r s i t y and i n t e gration. The c o n n e c t i o n s  i n aj|wilderness e c o s y s t e m a r e r e p r e s e n t e d by the  s t r u c t u r a l l i n k a g e s which p e r m i t i n f o r m a t i o n f l o w and m a t e r i a l c y c l i n g from one t r o p h i c l e v e l to a n o t h e r i n the community.  The i m p o r t a n t c o n n e c t i o n s  t h o s e l i n k s among s p e c i e s w h i c h d e f i n e the major dynamics o f the system.  are A  p r o m i n e n t example, which I l a t e r e x p l o r e a t l e n g t h , i s the l a r g e mammal p r e d a t o r - p r e y systems o f the k i n d found i n n o r t h e r n B r i t i s h C o l u m b i a .  A l s o of  paramount i m p o r t a n c e , tat.  i s the interdependence  between a s p e c i e s and i t s h a b i -  W i l d a n i m a l s a r e d e p e n d e n t upon t h e n a t u r a l environment  t o p r o v i d e ,food,  s p a c e , s h e l t e r , and p r o t e c t i o n . Management p o l i c i e s t h a t i g n o r e t h e s e key r e l a t i o n s h i p s a r e prone t o f a i l u r e o v e r t h e l o n g term.  There i s a growing  l i t e r a t u r e o f management f a i l u r e b e c a u s e t h e o r d e r o f t h e managed system was not u n d e r s t o o d  2.3  ( S m i t h , 1968; H a r r i s o n , 1965; Smith & Van den Bosch,  1967).  N o n - l i n e a r i t i e s and M u l t i p l e E q u i l i b r i a The r e l a t i o n s h i p between d e p e n d e n t and independent  e c o l o g i c a l systems i s o f t e n n o n - l i n e a r ( G r o s s , 1972).  variables i n  A prominent  example  i s t h e r e l a t i o n s h i p between p o p u l a t i o n d e n s i t y and p o p u l a t i o n growth.  A t low  p o p u l a t i o n d e n s i t i e s , p o p u l a t i o n growth i s u s u a l l y much f a s t e r than a t h i g h d e n s i t i e s where f o o d a v a i l a b i l i t y o r space may be a l i m i t i n g f a c t o r . T h e s e i n t r i n s i c n o n - l i n e a r r e l a t i o n s h i p s between b i o l o g i c a l v a r i a b l e s can sometimes r e s u l t i n t h e f o r m a t i o n o f e q u i l i b r i u m c o n d i t i o n s i n t h e p a r t i c u l a r system b e i n g c o n s i d e r e d .  A c l a s s i c example, and one t h a t i s im-  portant, t o t h i s a n a l y s i s , i s t h e s i m p l e r e c r u i t m e n t model d e r i v e d by p l o t t i n g p o p u l a t i o n m o r t a l i t y r a t e and r e p r o d u c t i v e r a t e a g a i n s t p o p u l a t i o n d e n s i t y . The growth o r d e c l i m e o f a p o p u l a t i o n i s d e t e r m i n e d b i r t h s and deaths i n t h e p o p u l a t i o n .  by t h e d i f f e r e n c e between  I f t h e m o r t a l i t y r a t e exceeds t h e b i r t h -  r a t e then t h e p o p u l a t i o n w i l l d e c l i n e ; w i t h r e p r o d u c t i o n g r e a t e r than mort a l i t y the population w i l l increase.  F o r prey s p e c i e s p r e d a t i o n i s c l e a r l y  one o f t h e n a t u r a l m o r t a l i t y f a c t o r s t h a t i n f l u e n c e s p o p u l a t i o n s t r e n d s . R e p r o d u c t i o n and m o r t a l i t y i n some n a t u r a l p o p u l a t i o n s a r e r e l a t e d to p o p u l a t i o n d e n s i t y , p a r t i c u l a r l y i n l a r g e , l o n g - l i v e d s p e c i e s w i t h low reproductive rates.  A t low d e n s i t i e s r e p r o d u c t i o n exceeds m o r t a l i t y which  a l l o w s t h e p o p u l a t i o n t o i n c r e a s e toward t h e c a r r y i n g c a p a c i t y o f t h e range.  14  reproduction  mortality  population  density  Figure 2.1: Density dependent population dynamics, r a f t e r Gross, 1972)  Figure 2 . 2 : Simple recruitment curve (modified after Gross, 1972)  15  However, when d e n s i t i e s reach c e r t a i n l i m i t s b i r t h rates decline with a concomitant or subsequent increase in m o r t a l i t y rates due to food shortage or crowding.  Presumably an e q u i l i b r i u m i s reached at V ,  the carrying capacity  of the h a b i t a t where the population i s more or less s t a b l e .  This i s the  generally accepted theory and i s usually conceptualized by w i l d l i f e managers (Gross, 1972) as shown i n Figure 2.1. The population density in any one year i s determined in large part by the density i n the preceding one; i . e . by the ' h i s t o r i c a l s e n s i t i v i t y ' of the system. The r e p r o d u c t i v i t y and m o r t a l i t y curves may be combined in a funct i o n which r e l a t e s any two successive populations as shown in Figure  2.2.  For any stock density less than k the death rate i s less than the reproductive rate and the population in the succeeding year t+1 w i l l increase by the amount  (Nt  +i-Nt)  as shown.  This increase represents the net annual growth  of the population and i s known as the recruitment or net annual increment. While Figures 2.1 and 2.2 i l l u s t r a t e the presence of a stable e q u i l i b r i u m in the system, i t should be noted that not a l l e q u i l i b r i a are stable.  Some are unstable or ' t r a n s i e n t ' and represent points at which the  system undergoes a dramatic s h i f t into a new configuration or else to extinction.  These unstable e q u i l i b r i a can i n d i c a t e c r i t i c a l thresholds which  have important i m p l i c a t i o n s f o r the i n t e g r i t y or s i z e of natural populations. S t a b i l i t y i s usually conceptualized as the property of a system to return to some e q u i l i b r i u m or 'ground p o s i t i o n '  ( H o l l i n g , 1973).  When a  system i s perturbed to a small or moderate degree i t tends toward recovery of i t s i n i t i a l state once the perturbation i s removed.  Systems variables  operating w i t h i n the l i m i t s of t h e i r h i s t o r i c a l experience, then, tend to o s c i l l a t e around an e q u i l i b r i u m point or, r a t h e r , w i t h i n a domain defined by certain s t a b i l i t y l i m i t s .  16  A s y s t e m p e r t u r b e d beyond t h e l i m i t s o f s t a b i l i t y , even i f o n l y by an i n c r e m e n t a l c h a n g e i n one o f t h e d r i v i n g v a r i a b l e s , can s h i f t a b r u p t l y and u n p r e d i c t a b l y f r o m one s t a b i l i t y domain c h a n g e may  be i r r e v e r s i b l e .  to another ( H o l l i n g , 1978).  Situations i n which ecosystem s t a b i l i t y  have been o b s e r v e d have been documented ies  son,  shifts  f o r the Great Lakes commercial  ( B e e t o n , 1969; C h r i s t i e , 1974), a l g a l  forest ecosystems  This  blooms  fisher-  i n Lake E r i e ( S h a p i r o , 1973),  (Gomez Pompu e t a l , 1972; N i e r i n g & G o o d w i n , 1974; H u t c h i n -  1970), i n s e c t p e s t p o p u l a t i o n s i n A s i a , Europe and N o r t h A m e r i c a  & K i r i t a n i , 1975; J o n e s , 1975; Southwood  and Comins,  1976;  (Saraba  I s a e v and K l e b o p r o s ,  1977), and g r a z i n g systems i n N o r t h A m e r i c a , A f r i c a and A u s t r a l i a ( G l e n d e n i n g , 1952; N o y - M e i r , 1 9 7 5 ) .  Such phenomena c l e a r l y have i m p o r t a n t i m p l i c a t i o n s f o r  managers i n s i t u a t i o n s where the components  o f a n a t u r a l s y s t e m , s u c h as a  w i l d l i f e p o p u l a t i o n , a r e b e i n g m a n i p u l a t e d through management Our knowledge know t h e c r i t i c a l  practices.  o f e c o s y s t e m s i s s u c h t h a t i n m o s t c a s e s we d o n o t  t h r e s h o l d s o f the p a r t i c u l a r system being managed.  Shifts  in system behaviour g e n e r a l l y remain u n p r e d i c t a b l e . If a w i l d e r n e s s ecosystem s u c h a s t h e S p a t s i z i w e r e t o b e l e f t e n t i r e l y a l o n e ( i . e . no_ h u n t i n g a n d  fishing  o r o t h e r h u m a n d i s t u r b a n c e ) t h e n t h e r e w o u l d b e no n e e d t o u n d e r s t a n d i t s d y namics.  I t has e v o l v e d and p e r s i s t e d f o r t h o u s a n d s o f y e a r s n a t u r a l l y  t h e r e a p p e a r s n o r e a s o n why  i t c o u l d n o t c o n t i n u e t o do s o .  hand, t h e e c o s y s t e m i s t o be e x p l o i t e d critical  I f , on t h e o t h e r  t o some e x t e n t t h e n t h e d e f i n i t i o n  of  t h r e s h o l d s becomes a p r i o r i t y f o r management c o n c e r n ; t h a t i s , i f  e x p l o i t a t i o n i s not t o impede  2.4  and  Dynamic  natural  processes nor impact natural  Variability  Natural systems are in a continual ecological  communities.  s t a t e of change.  Variation in  p a r a m e t e r s s u c h as p o p u l a t i o n numbers, s p e c i e s c o m p o s i t i o n , e q u i l i -  b r i u m c o n d i t i o n s and so f o r t h shapes t h e p a t t e r n o f e c o s y s t e m p r o g r e s s i o n and  succession.  The 'pathways' o f t h e s e v a r i a b l e s o v e r time r e p r e s e n t s t a b l e  t r a j e c t o r i e s ( i . e . s t a b l e v a r i a t i o n a b o u t an e v o l v i n g o r c h a n g i n g mean).  Dy-  namic change can be g e n e r a t e d w i t h i n t h e system as a consequence o f i n t r i n s i c t r i g g e r i n g mechanisms and f e e d b a c k l o o p s o r e x o g e n o u s l y as a consequence o f u n p r e d i c t e d , random e v e n t s .  D e n s i t y dependent phenomena between competing  s p e c i e s , o r between p r e d a t o r s and t h e i r p r e y , a r e examples o f the f o r m e r . S t o c h a s t i c e v e n t s such as f o r e s t f i r e s , f l o o d s , c o l d s n a p s , and a c c i d e n t s r e p r e s e n t e x t e r n a l l y imposed f o r c e s f o r change i n e c o l o g i c a l systems.  Such  e v e n t s produce a dynamic v a r i a b i l i t y which i s an i m p o r t a n t c o n t r i b u t o r t o t h e s t a b i l i t y of ecosystems.  The system's c a p a c i t y t o r e s p o n d t o and a b s o r b  change i s a measure o f i t s r e s i l i e n c e .  The w i d e r the range o f v a r i a b i l i t y  ( o f e x p e c t e d and u n e x p e c t e d change) t h e system can accommodate, the more r e s i l i e n t i t i s and, hence, t h e g r e a t e r i t s o p p o r t u n i t i e s f o r s u r v i v a l .  As  H o l l i n g (1978) n o t e s , management p o l i c i e s t h a t r e d u c e v a r i a b i l i t y i n time and space tend t o l e s s e n t h e r e s i l i e n c e o f t h e system ,and,hence, a r e q u e s t i o n a b l e i n terms o f t h e i r e c o l o g i c a l s o u n d n e s s .  I t i s the h i s t o r i c a l e x p e r i e n c e o f a  s y s t e m , o r sub-system such as a s p e c i e s p o p u l a t i o n , which d e t e r m i n e s i t s o v e r all resilience.  An h i s t o r i c a l ' s e n s i t i v i t y ' i s so d e v e l o p e d t h r o u g h t i m e  which a t t u n e s t h e system f o r f u t u r e e v e n t s ( H o l l i n g , 1968). 2. 5 U n c e r t a i n t y The i s s u e o f u n c e r t a i n t y i n e c o l o g i c a l management has been r a i s e d earlier.  N e v e r t h e l e s s , i t w a r r a n t s f u r t h e r d i s c u s s i o n here as i t i s c e n t r a l  to our u n d e r s t a n d i n g o f b i o l o g i c a l systems.  The t o t a l of what we know, what  i s f a c t about n a t u r a l e n v i r o n m e n t s and., p a r t i c u l a r l y , w i l d l i f e r e s o u r c e s p a l e s b e s i d e t h e sum o f our i g n o r a n c e .  How t o d e a l w i t h the unknown and, i n -  deed, how t o b e n e f i t from i t where p o s s i b l e i s u n d o u b t e d l y an i m p o r t a n t conc e r n f o r management.  I t becomes i n c r e a s i n g l y i m p o r t a n t i n s i t u a t i o n s where  18 management is interventionist or manipulative in character. This is the case with Spatsizi where manipulative actions are allowed in the form of hunting and fishing.  Here, uncertainties arise principally  in the determination and regulation of harvests.  Substantial sources of error  can be introduced at the data gathering stage or during analysis.  Fundamental  assumptions regarding the dynamics or structure of the system may be erroneous.  However, whatever the nature of the uncertainty the intent of the Order-  in-Council is clear with respect to the obligation placed on park managers. Management strategies must incorporate some mechanism of reducing uncertainties to the greatest extent possible i f harvests are to be sustained without negatively influencing the natural balance of the wildlife community.  C H A P T E R  III  Predator-Prey Systems  3.1  Introduction The presence of predator species in a w i l d l i f e community represents  an a d d i t i o n a l t r o p h i c l e v e l i n the ecosystem and, therefore, higher complexity than in those communities lacking predators.  In most northern temperate en-  vironments the large t e r r e s t r i a l predators such as the bear, wolf, and cougar have been l a r g e l y eliminated due to competition f o r prey and habitat by man. No s i g n i f i c a n t w i l d populations of large predators, for instance, remain in western Europe or the Scandinavian c o u n t r i e s .  In North America such popula-  tions e x i s t as s m a l l , protected refugia in conservancy areas; remote and usua l l y dwindling unprotected enclaves in the south; and as widely but sparsely d i s t r i b u t e d populations in the large t r a c t s of wilderness that remain in the Canadian north and Alaska.  In northern B r i t i s h Columbia, including the Spat-  s i z i , one of the most d i s t i n c t i v e features of the wilderness ecosystem are i n t a c t , large mammal predator-prey systems.  20 H i s t o r i c a l l y , man  has c o m p e t e d f o r s p e c i e s t h a t c o m p r i s e  prey f o r  other predators.  Of p a r t i c u l a r n o t e a r e l a r g e u n g . u l a t e s , some o f w h i c h  have  been domesticated  to a i d i n the development of a g r i c u l t u r a l  Such  societies.  c o m p e t i t i o n h a s t y p i c a l l y p r o m p t e d a c o n c e r t e d a c t i o n o n t h e p a r t o f h u m a n .• b e i n g s , i n d i v i d u a l l y and c o l l e c t i v e l y , t o s u p p r e s s Since e a r l y times predator animal  husbandry.  'control'  or eliminate predators.  has been most i m p o r t a n t  More r e c e n t l y , the advent  in relation  of 'sport' hunting  in  s o c i e t i e s has s o m e t i m e s c r e a t e d a demand f o r c o n t r o l p r o g r a m s w h i c h duce the take of w i l d ungulates er  by p r e d a t o r s a n d ,  hence,  to  western would  allow increased  rehunt-  harvests. Predator control to 'aid' wild ungulates, s t i l l  some c i r c l e s , it is designed  widely promoted i n  i s a good e x a m p l e o f an i n t e r v e n t i o n i s t management s t r a t e g y , f o r to a c t i v e l y manipulate  a l t e r the order of the  i n t e r s p e c i f i c r e l a t i o n s h i p s and,  system.  I n t h i s s t u d y , my  c o n c e r n w i t h p r e d a t o r s as an i n t e g r a l c o m p o n e n t o f  the w i l d l i f e community i s t w o f o l d . r e n t l y p r a c t i c e d in S p a t s i z i Park,  F i r s t , while predator c o n t r o l i s not t h e r e have been r e c e n t p r o p o s a l s to  tute i t . Second, c e r t a i n predator species form part of the hunter area, a f a c t which  has c e r t a i n i m p l i c a t i o n s i n terms  ments being s a t i s f i e d .  sary s c i e n t i f i c information to s a f e l y enable  system?  ture of the w i l d l i f e community?  Do we  instiin the  however  have the  the h a r v e s t o f a few  i m p a i r i n g the i n t e g r i t y of the n a t u r a l system  kill  cur-  of the p o l i c y r e q u i r e -  What a r e t h e e f f e c t s o f p r e d a t o r r e m o v a l ,  s t r i n g e n t l y r e g u l a t e d , upon the p r e d a t o r - p r e y  without  hence,  neces-  predators  or a l t e r i n g the s t r u c -  F u r t h e r , e x a c t l y what kinds of i n f o r m a t i o n  w o u l d be r e q u i r e d t o m a k e t h i s d e c i s i o n ? I n t h i s c h a p t e r my . . o b j e c t i v e i s t o a p p l y s o m e o f t h e g e n e r a l cepts outlined in Chapter to  identify crucial  c o n s i s t e n t with the  II to the dynamics o f p r e d a t o r - p r e y  concerns  systems  conin order  f o r f o r m u l a t i n g management s t r a t e g i e s t h a t a r e  Order-in-Counci1.  21 I b e g i n w i t h a r e v i e w o f t h e t h e o r e t i c a l and e m p i r i c a l e v i d e n c e p e r t i n e n t to p r e d a t o r - p r e y systems  i n g e n e r a l , d e s c r i b i n g a model o f p r e d a t o r -  ungulate p o p u l a t i o n dynamics that i s r e p r e s e n t a t i v e of northern B r i t i s h Columbia.  3.2  P r e d a t o r - P r e y Dynamics The i n t e r a c t i o n b e t w e e n p r e d a t o r s and t h e i r p r e y has b e e n one o f  t h e m o s t p o o r l y u n d e r s t o o d and p r o b a b l y c o n t r o v e r s i a l a s p e c t s o f w i l d l i f e logy.  eco-  P r e d a t i o n i s v i e w e d by some a u t h o r i t i e s as a l i m i t i n g f a c t o r t o t h e  g r o w t h o f c e r t a i n p r e y p o p u l a t i o n s - an i d e a h o t l y c o n t e s t e d by o t h e r s . d a t o r - p r e y systems  Pre-  t r a d i t i o n a l l y h a v e b e e n s i m p l i s t i c a l l y c o n c e p t u a l i z e d by  w i l d l i f e m a n a g e r s , r e s u l t i n g commonly i n d i s a s t r o u s c o n s e q u e n c e s f o r t h e p o p u l a t i o n b e i n g m a n a g e d , o r unmanaged as t h e c a s e may  be.  P r e d a t o r s a r e d e p e n d e n t upon t h e a v a i l a b i l i t y o f o t h e r a n i m a l cies ( i n c l u d i n g , sometimes, other predators) f o r t h e i r food  spe-  requirements.  P o p u l a t i o n s o f p r e d a t o r s and t h e i r p r e y a r e i n c o n t i n u a l f l u c t u a t i o n .  Some-  t i m e s t h e c y c l e s a p p e a r t o be c a u s a l l y l i n k e d and a t o t h e r t i m e s t h e v a r i a t i o n i n one p o p u l a t i o n seems t o h a v e no c o r r e l a t i o n t o c h a n g e i n t h e o t h e r . dynamics of prey p o p u l a t i o n s , however, a p p a r e n t l y have q u a l i t a t i v e l y c h a r a c t e r i s t i c s i n communities without.  The  distinct  w i t h p r e d a t o r s p r e s e n t as o p p o s e d t o t h o s e  This warrants c l o s e r examination because of the e f f e c t s predator  h a r v e s t s may  h a v e on t h e r e m a i n i n g p r e y p o p u l a t i o n s .  I n C h a p t e r I I I u s e d t h e r e p r o d u c t i o n and m o r t a l i t y c u r v e s t o demons t r a t e t h e t h e o r e t i c a l e x i s t e n c e o f s y s t e m e q u i l i b r i u m ; i n t h i s c a s e 'k', c a r r y i n g c a p a c i t y of the range. however, are r a r e l y t h i s s i m p l e .  the  R e p r o d u c t i o n and m o r t a l i t y c u r v e s i n n a t u r e , R e c e n t e x p e r i m e n t a l r e s e a r c h and c o m p u t e r  s i m u l a t i o n s t u d i e s have demonstrated  t h a t the topography  c u r v e s i s much more c o m p l e x t h a n ' h i t h e r t o u n d e r s t o o d .  of prey recruitment In l a r g e p a r t , t h i s  a r i s e s because the response of p r e d a t o r s to v a r y i n g prey d e n s i t i e s appears  to/  22 be n o n - l i n e a r .  E s c a p e a b i l i t y o f the prey and the f a i l u r e o f p r e d a t o r s to form  s e a r c h images f o r r a r e prey c o n s t i t u t e a f u n c t i o n a l r e s p o n s e i n p r e d a t o r s t h a t d i m i n i s h e s t h e i r impact upon prey  m o r t a l i t y a t v e r y low prey d e n s i t i e s . As  prey d e n s i t i e s i n c r e a s e t h e r e i s a d e n s i t y d e p e n d e n t i n c r e a s e i n prey m o r t a l i t y as t h e s e e f f e c t s a r e overcome, f o l l o w e d by an i n v e r s e d e n s i t y dependent limb i n d u c e d by s a t i a t i o n o f the p r e d a t o r and a r e d u c e d n u m e r i c a l r e s p o n s e ( H o l l i n g , 1973).  The net r e s u l t i s the e x i s t e n c e o f a p r e d a t o r - i n d u c e d 'bulge' i n the  prey m o r t a l i t y c u r v e a t low-moderate d e n s i t i e s ( R i c k l e f s , 1976). The r e p r o d u c t i o n c u r v e f o r prey i s a l s o d i v e r g e n t a t v e r y low densities.  R e p r o d u c t i o n r a t e s a r e l i k e l y t o p l u n g e s h a r p l y , perhaps a p p r o a c h i n g  z e r o , as d i f f i c u l t i e s i n f i n d i n g mates i m p a i r s b r e e d i n g . The r e s u l t o f n o n - 1 i n e a r . d e n s i t y d e p e n d e n t e f f e c t s a t low prey dens i t i e s a l t e r s the c o n v e n t i o n a l c o n c e p t u a l i z a t i o n o f the r e p r o d u c t i v i t y / m o r t a l i t y r e l a t i o n s h i p as d e p i c t e d by G r o s s (1972) ( F i g u r e 3.1). Combining t h e s e two c u r v e s p r o d u c e s a r e c r u i t m e n t c u r v e f o r prey s p e c i e s as shown i n F i g u r e 3.2.  Note t h a t on t h i s r e f i n e d c u r v e t h e r e a r e  'now f o u r e q u i l i b r i u m p o i n t s ; two s t a b l e e q u i l i b r i a (E ) S  k and two t r a n s i e n t o r u n s t a b l e e q u i l i b r i a (f; ) t  a t p o p u l a t i o n s b and  a t a and c.  Under optimum  c o n d i t i o n s prey p o p u l a t i o n s a p p a r e n t l y tend to grow towards the c a r r y i n g capac i t y k where d e n s i t y dependence i s the p r i n c i p a l l i m i t i n g f a c t o r . I f , f o r any number o f r e a s o n s , p o p u l a t i o n s a r e d e p r e s s e d to the u n s t a b l e e q u i l i b r i u m l e v e l at a then t h e y w i l l c o n t i n u e t o d e c l i n e t o the low e q u i l i b r i u m d e n s i t y a t b.  Below the low e q u i l i b r i u m l i e s a n o t h e r u n s t a b l e e q u i l i b r i u m a t a from  which p o i n t p o p u l a t i o n s w i l l d e c l i n e to e x t i n c t i o n . F u r t h e r e l a b o r a t i o n on p r e d a t o r - p r e y e q u i 1 i b r i u m models i s g i v e n by R i c k l e f s (1976). What k i n d o f e m p i r i c a l s u p p o r t i s t h e r e f o r such a model o f p r e d a t o r prey dynamics?  The e v i d e n c e i n d i c a t e s t h a t a r e m a r k a b l e s i m i l a r i t y i n the  t o p o l o g y o f prey r e c r u i t m e n t p a t t e r n s e x i s t s not o n l y w i t h i n insect/mammal systems where much o f the o r i g i n a l work was done ( H o l l i n g , 1966 &  1976;  a  b  e  population  density  Figure 3.1: Density dependent population dynamic (modified after Holling, 1973)  E  -• stable equilibrium  Et  - transient  Population in year t+1  population  Figure 3.2:  in  year  t  Prey species recruitment curve.  24 M a c l e o d , 1978) b u t a l s o e x t e n d s i n t o f i s h e r i e s ( R i c k e r , 1954; H o l l i n g , 1973; Peterman & G a t t o , 1977), s m a l l mammal systems (Brand 1976 e t a l ; P e a r s o n , 1966) and w o l f - c a r i b o u , wolf-moose  systems (Haber e t a l , 1976; Haber, 1977; W a l t e r s  e t a l , 1978). Moreover, there i s evidence t o i n d i c a t e t h a t depressed p o p u l a t i o n s can be m a i n t a i n e d i n low d e n s i t y s t a b i l i t y domains even i f t h e c a u s a l f a c t o r t h a t i n i t i a t e d t h e s h i f t i s removed. versible.  In o t h e r words, t h i s s h i f t may be i r r e -  In t h e c a s e o f t h e G r e a t Lakes F i s h e r i e s , Smith (1968) and Beeton  (1969) document p r e c i p i t o u s d e c l i n e s i n l a k e t r o u t , h e r r i n g , and w h i t e f i s h immediately f o l l o w i n g years o f sustained high harvests. S i m i l a r l y , c o n t r o l of p e r c h i n Lake Windermere t o enhance s a l m o n i d s t o c k s s e v e r e l y r e d u c e d t h e p o p u l a t i o n s t o such an e x t e n t t h a t a l t h o u g h c o n t r o l c e a s e d i n 1947 t h i s s p e c i e s has shown no t e n d e n c y t o r e t u r n t o o r i g i n a l l e v e l s (Le Cren e t a l , 197-2). H o l l i n g (1973) a r g u e s , "whatever t h e s p e c i f i c c a u s e s , i t i s c l e a r t h a t t h e p r e c o n d i t i o n f o r t h e c o l l a p s e was s e t by t h e h a r v e s t i n g o f f i s h . , even though d u r i n g a l o n g p e r i o d t h e r e were no o b v i o u s s i g n s o f problems.  The f i s h i n g  a c t i v i t y , however, p r o g r e s s i v e l y r e d u c e d t h e r e s i l i e n c e o f t h e system so t h a t when t h e i n e v i t a b l e u n e x p e c t e d e v e n t o c c u r r e d , t h e p o p u l a t i o n s c o l l a p s e d . " Haber (1977) s u g g e s t s a s i m i l a r phenomenon f o r c a r i b o u herds i n A l a s k a and the Yukon.  These p o p u l a t i o n s , he a r g u e s , have been p e r i o d i c a l l y m a i n t a i n e d  at low d e n s i t i e s due t o s e v e r e h u n t i n g p r e s s u r e . The q u e s t i o n might r e a s o n a b l y be asked what d i f f e r e n c e does i t make i f a p a r t i c u l a r s p e c i e s p o p u l a t i o n i s m a i n t a i n e d i n a low d e n s i t y s t a b i l i t y domain as opposed t o a h i g h one?  In t h e c a s e o f S p a t s i z i would a s h i f t from  h i g h t o low s t a b l e p o p u l a t i o n s o f c a r i b o u , f o r i n s t a n c e , v i o l a t e t h e c o n d i t i o n s s e t o u t by t h e O r d e r - i n - C o u n c i 1 ? Some p o s s i b l e e f f e c t s c a n e a s i l y be dis.cerned. F i r s t , t h e r e may be o t h e r i m p o r t a n t b u t unknown d e n s i t y dependent f a c t o r s which w i l l a f f e c t popul a t i o n s a t low d e n s i t i e s b u t n o t h i g h e r ones.  Second, i t might then be p o s -  - s i b l e f o r predators to f u r t h e r reduce prey populations to the point of extinction.  T h i r d , u n d e r - u t i 1 i z a t i o n of the habitat by the reduced species may  r e s u l t in s t r u c t u r a l changes i n the vegetation and/or animal community.  All  these, plus other unpredictable f a c t o r s , could r e s u l t in i r r e v e r s i b l e change i n the natural systems and communities of the area.  Such change would contra-  vene the i n t e n t of the p o l i c y mandate.  3.3  Predator Response and A l t e r n a t e Prey One f i n a l point remains.  As with i n d i v i d u a l species  populations,  the r e l a t i v e abundance of predators and prey at a given time, t h e r e f o r e , i s also l a r g e l y determined by the h i s t o r i c a l s e n s i t i v i t y or 'memory' of the system.  A steady-state i s never reached because of the lag e f f e c t s and the f a c t  that there always e x i s t s a 'wrong' number of predators f o r any prey population ( c f . Figure 3.3).  instantaneous  In any one generation t h i s 'memory'  is  modified by environmental f a c t o r s and density dependent r e l a t i o n s h i p s between prey and predators.  These modifications r e s u l t in s p a t i a l v a r i a b i l i t y which  determines the extent to which any perturbation w i l l spread throughout the system.  When v a r i a b i l i t y in s p a t i a l and h i s t o r i c a l s e n s i t i v i t y is high the  system i s more s t a b l e . The recruitment curve provides a means of examining the e f f e c t of stochastic v a r i a b i l i t y upon population dynamics.  Figure 3.4 i l l u s t r a t e s the  range of v a r i a t i o n in recruitment that a given population, N* , might y i e l d . Conversely, note that determination of a recruitment l e v e l in a f i e l d survey does not. i n f e r a p a r t i c u l a r population density but, r a t h e r , a range of densities.  This has important i m p l i c a t i o n s in s i t u a t i o n s where management b i o -  l o g i s t s e s t a b l i s h harvest l e v e l s f o r a population based s o l e l y on recruitment data. The response of predators to changes in prey populations is of two basic kinds ( R i c k l e f s 1976).  The short-term, or f u n c t i o n a l response, i s simply  26  Figure 3.3: Time lags in predator-prey systems (source R i c k l e f s , 1976, p.257 & 308)  Population Density in year t+i  N*  Population Density in year t  Figure 3.4: Effects of environmental variability on recruitment where solid line represents the theoretical recruitment curve; dashed lines represent maximum and minimum recruitment curves under benign and adverse environmental conditions respectively; a - c represents, therefore, the spread of environmental, density independent effects at. population N*; a-d represents maximum possible recruitment 9 N*,• c-d represents minimum recruitment •@ N*.  the resulting change in the number of prey killed and eaten over a specified time interval (i.e. attack rate).  The number of prey taken by an individual  predator is a function of prey density, the predator's attack rate, and the time spent searching.  The attack rate, in turn, is dependent upon the pre-  dator's searching efficiency and the effective exposure time to prey. The time a predator spends searching for prey equals total time minus the 'handling' time - the time spent in the actual k i l l and consumption of prey prior to the next search.  While handling time appears to be a constant there  is usually a learning function associated with each predator that increases the sarching efficiency and, hence, the attack rate.  This increases rapidly  at f i r s t and then its effect diminishes with each additional k i l l made. In addition, a predator tends to eat more prey at higher densities but does so at a decelerating rate so that an inversely density dependent response relationship exists.  Numerical and functional responses sometimes interact since  a change in attack rate of an individual predator may be dependent upon predator density. The long-term, or numerical response, is a decline in the number of predators triggered by a continuing decline in prey.  Similarly, an in-  crease in prey can generate an increase in predators. This numerical response hinges upon the nutritional theory of the predator parent as i t is exposed to varying prey densities.  Not only the  number of progeny are affected, but also the quality as reflected in their health, size, vigour, and attack efficiency (Moiling, 1968).  Predator popu-  lations generally respond to changes in prey populations by either increasing or congregating in areas of high prey density.  Such 'tactics' can occur  jointly or separately during the natural regulation of prey numbers (Readshaw, 1973). In many instances, particularly in mammal predator-prey systems, the  food of predators consists of a number of alternative prey species, some more preferred than others.  The relative frequency of a particular prey eaten is  a function of the relative density of prey in the environment modified by the predator's preference for one over the other (Murdoch, 1973).  A stabilizing  mortality effect can be produced on less-preferred prey populations at the times when they are on the increase concomitant with decreasing preferred prey. Furthermore, prey preference can vary as concentrations of alternative prey change with respect to each other. As preferred prey populations diminish predators will switch their searching efforts to more abundant alternatives.  Thus, the availability of  alternate sources of food can maintain predator populations at levels higher than would have resulted i f only a single prey species existed. One of the obvious effects of predator 'switching' is to reduce the impact upon a preferred species population decline. On the other hand, with alternate prey "subsidizing" high predator levels then any attempt at population rebound by the depleted preferred species may not be successful because of the disproportionately high number of predators.  In the extreme case i t may be that  sustained high predator density can accelerate the decline of a preferred species.  Shaw (1977 a & b) concluded that increased impact upon declining  deer populations resulted from high mountain lion densities sustained by livestock.  Similarly, Peterson (1976) and Mech & Karns (1977) observed that  the availability of beaver actually increased the wolf population during a moose decline which resulted in an accelerated mortality for the moose. What is the nature of the field evidence on record relating to the interrelationship between predators'and-ungulates?  Before reviewing the  evidence i t is worthwhile to note that there are three principal views held by wildlife managers concerning the effects of predators upon prey species.. The f i r s t is that predators do act as a limiting factor in controlling prey  30 populations. Still  Others h o l d t h a t p r e d a t i o n has no c o n t r o l l i n g e f f e c t upon p r e y .  o t h e r s b e l i e v e t h a t p r e d a t o r s s e r v e t o i n c r e a s e prey p o p u l a t i o n .  Such  d i v e r g e n c e o f o p i n i o n has sparked f r e q u e n t and b i t t e r c o n t r o v e r s y i n r e c e n t y e a r s both i n the U.S.A. and Canada about how w i l d u n g u l a t e p o p u l a t i o n s  can  and s h o u l d be managed. The answer l i k e l y l i e s i n the f a c t t h a t p r e d a t o r s can e x e r t a l l o f these e f f e c t s s e p a r a t e l y upon prey under d i f f e r e n t c o n d i t i o n s depending the  upon  h i s t o r i c a l s e n s i t i v i t y o f the p a r t i c u l a r system and the key e n v i r o n m e n t a l  i n f l u e n c e s a c t i n g on the system d u r i n g any s p e c i f i c time i n t e r v a l .  Resolution  of the c o n t r o v e r s y becomes c l e a r e r i f the e m p i r i c a l e v i d e n c e d e r i v e d from f i e l d experience i s  examined.  P r e d a t i o n , of c o u r s e , i s one o f s e v e r a l agents  t h a t c o n t r i b u t e to  the n a t u r a l m o r t a l i t y r a t e s of those s p e c i e s t h a t s e r v e as food t o o t h e r s . If  p r e d a t o r s are s e l e c t i v e i n t h e i r c a p t u r e and k i l l - o n l y the weak,  infirm,  or d i s e a s e d i n d i v i d u a l s t h a t o t h e r w i s e are about t o succumb t o a n a t u r a l d e a t h , t h e n , c l e a r l y , they have a s u p p l a n t i n g o r "compensatory" the prey p o p u l a t i o n . dation  'compensates'  Compensatory  e f f e c t upon  p r e d a t i o n means s i m p l y t h a t death by p r e -  f o r , or r e p l a c e s , death t h a t was o t h e r w i s e imminent due  to o t h e r n a t u r a l causes and, t h e r e f o r e , has no e f f e c t upon the b r e e d i n g p o t e n tial  o f the p o p u l a t i o n .  If,  on the o t h e r hand, they remove v i g o r o u s  h e a l t h y , i n d i v i d u a l s which are s t i l l  p a r t o f the b r e e d i n g s t o c k , then they  have an a d d i t i v e . o r - "non-compensatory" Compensatory  and  effect.  p r e d a t i o n would r e q u i r e t h a t p r e d a t o r s have some means  of s e l e c t i n g i n d i v i d u a l s from c e r t a i n c a t e g o r i e s w i t h i n the p o p u l a t i o n .  There  i s good e v i d e n c e t h a t c a n i d p r e d a t o r s do s e l e c t s i c k , d i s e a s e d , or weak animals from a hunted p o p u l a t i o n .  Murie  (1940)  observed p r e f e r e n t i a l s e l e c t i o n of  heal thy-mule deer by coyotes and of d i s e a s e d Da 11 sheep by wolves  1946):  Mech (1966) and P e t e r s o n  (1976)  (Murie,  of moose by w o l v e s ; Ozo.ga and Harger  less  (1966) of white-tailed deer by coyotes; and Miller (1975) of caribou by wolves.  Pimlott (1967) observes that predation tends to be concentrated on  both young and old individuals and the predation on the former is compensatory i f i t is restricted to individuals which would likely have succumbed to a natural death anyway.  It seems reasonable to expect that incapacitation or  sickness in prey animals increases their vulnerability to predators.  Mech  (1970) has found, for example, that moose infested with tapeworms suffer increased predation because of their diminished strength in escaping from or warding off wolves. On the other hand, some authorities argue that the health of prey animals does not influence their vulnerability to predation.  Stephenson and  Johnson (1973) and Franzmann and Arneson (1976) have observed  non-discrimina-  tory attacks on moose by wolves; Neil sen (1975) observed similar patterns with coyotes preying on mule deer young and Beale and Smith (1973) found that bobcats were random in their attacks on pronghorn fawns.  It could be argued,  however, that attacks on healthy young do represent selective, although noncompensatory, predation as young constitute a generally more vulnerable class in the population.  More importantly though, i t may be that the observation of  non-discriminatory attacks is the wrong criterion by which to measure predator selectivity.  While attacks may be non-discriminatory, the degree of success  is likely to be directly correlated with the health and vigour of the prey. In other words, attacks on weak, young and infirm animals are probably more successful than attacks on healthy prey.  The end result is that while preda-  tors may not actively search out weak or diseased animals, healthy animals are better able to withstand attacks and thus enjoy a higher rate of escape. Another problem with many studies of this kind is that i t is usually d i f f i c u l t to rule out other causative mortality agents or to detect some factor that might be acting as an environmental the animal's susceptibility to attack.  stressor and would contribute to  For example, while field investigation  32  have shown that wolves eat young animals they do not adequately demonstrate the predator was responsible for the death of the individual.  After a l l ,  there is ample evidence indicating significant, natural neo-natal mortality of young ungulates in areas where predators are extirpated. An indirect effect of compensatory predation is its apparent capacity to stimulate reproductive rates in prey population.  Mech (1966, 1970),  for example, observed an increase in moose twinning from 6% to 38% which he concluded was a short-term effect of predation.  Other environmental stressors  and density dependent phenomena such as deteriorating-range can accentuate the impact of predation by increasing natural mortality or diminishing fecundity of the prey populations (Pimlott, 1967). It appears that most ungulates can sustain some level of predation without overall impairment to the health and numbers of the population.  Above  this level, predation can exert non-compensatory effects and thus can become an important mortality agent.  If wolf densities are high then the non-compen-  satory component of predation may become a significant limiting factor on prey populations (Pimlott, 1967). Conclusions drawn from field experience in North America suggest that predators generally do not acquire the capability of limiting the densities of prey populations in natural systems.  In the case of ungulates, seldom have  predators been implicated as the sole cause of population decline. Sometimes predators can act as a "carrying capacity adjuster" by amplifying prey mortality in situations of increased environmental stress (e.g. severe winters). In other instances, they can impede recovery of ungulate densities that have declined due to other causes and can accelerate declines. In the foregoing sections I have attempted to describe some of the important aspects of predator-prey systems that have management implications for populations which are to be harvested while maintaining the 'integrity'  33  of the wildlife community.  The f i r s t key principle to be noted is that the  relationship between predator and prey densities is non-linear so that at certain densities predation effects are pronounced while at others they may be negligible.  Thus, information pertaining to both predator and prey den-  sities is crucial for managing the harvest of either population under a preservationist mandate. Second, the system is ordered in that predator response may be geared to the relative abundances of alternate prey species.  Where alternate prey are  a significant element in the wildlife community the regulation of harvest activity must anticipate possible effects to other populations besides the hunted one(s). Third, predator-prey systems may be characterized by multiple' equilibria with at least two stability domains.  If maintenance of the prevailing  natural equilibrium state of wildlife communities is a policy requirement, then hunter harvests must be regulated so that critical thresholds are not transgressed. Finally, the effects of random, stochastic change introduce considerable unpredictability in the dynamics of wildlife populations.  Managers will  never have complete and totally accurate knowledge of the ecological system they are managing.  Because irreversibility can characterize these systems  the application of consumptive or exploitative policies must be done with great care.  34  CHAPTER  IV  The Effects of Hunting  4.1  Introduction One of the fundamental doctrines of contemporary wildlife. manage-  ment is that game populations can be harvested at rates not exceeding the net annual increment and s t i l l maintain a flourishing breeding stock. As illustrated in Chapter III, the maximum recruitment rate of any population is reached at some density below the carrying capacity (Ricker, 1954; Beverton and Holt, 1957; Watt, 1968).  Since recruitment is a measure of the overall  growth of the population as defined by the difference in intrinsic mortality and fecundity rates of the population, i t theoretically represents, therefore, a biological surplus of animals that can be removed while maintaining a stable population.  Technically speaking, recruitment, measures the addition of young  to the breeding stock once neo-natal mortality losses have been accounted for. Theoretically, i t also includes incremental losses or gains through immigration or dispersal.  In the case of large mammals in northern mountain habi-  tats recruitment should usually be measured after the f i r s t winter; i.e. the percentage of yearlings in tne population (Haber, 1978; Bergerud, 1978a). In setting harvest levels for each species, managers have generally lacked information in two vital areas.  First i t has almost always been im-  35 p o s s i b l e t o s e c u r e t h e d a t a n e e d e d f o r an a c c u r a t e d e t e r m i n a t i o n o f t h e biological  surplus of the harvested  populations.  Second, the e f f e c t s of  hunting upon a w i l d l i f e p o p u l a t i o n a r e n e i t h e r q u a l i t a t i v e l y nor q u a n t i t a t i v e l y well understood.  T h e i m p l i c a t i o n o f b o t h t h e s e p o i n t s i s t h a t b i g game h a r -  vests have not been w e l l r e g u l a t e d i n the past. t y p i c a l l y gone u n r e g u l a t e d  until  have  i t has been p e r c e i v e d t h a t a p a r t i c u l a r popu-  l a t i o n was b e i n g s i g n i f i c a n t l y d e p l e t e d t h r o u g h instances, the suspension  In f a c t , h a r v e s t s  overkill.  Sometimes  o r s e v e r e r e s t r i c t i o n o f h u n t i n g has been  i n such sufficient  t o o f f s e t f u r t h e r p o p u l a t i o n d e c l i n e s , i n o t h e r s , c o r r e c t i v e r e g u l a t i o n comes too l a t e and l o c a l  populations are extirpated.  U t i l i t a r i a n use i s a p r i o r i t y management B.C.'s b i g game p o p u l a t i o n s .  However,  p r e s e r v a t i o n i s t h e key management gers to ensure  prudent  i n an a r e a such as t h e S p a t s i z i where  concept,  t h e o b l i g a t i o n o f w i l d l i f e mana-  harvest strategies i s greatly increased.  faire" approach to conventional t e c t i o n i s t management  o b j e c t i v e f o r most o f  The  h a r v e s t r e g u l a t i o n must be a b a n d o n e d .  u n l e s s due t o n a t u r a l c a u s e s .  Spatsizi  4.2  Pro-  a c t i o n m u s t be a n t i c i p a t o r y r a t h e r t h a n r e a c t i v e .  p l e t i o n o f w i l d l i f e p o p u l a t i o n s m u s t n o t be a l l o w e d t o o c c u r t o a n y  ments f o r management  "laissez-  What, t h e n , comprises  De-  extent,  the information require-  p r e s c r i p t i o n s to achieve the stated objectives f o r  Park?  •Quantitative Aspects  of Maximizing  Harvests  The f i r s t c a t e g o r y o f i n f o r m a t i o n r e l a t e s to the d e t e r m i n a t i o n biological  or "harvestable" surplus.  F i g u r e 4.1  of  embellishes the c h a r a c t e r i s -  t i c m u l t i - e q u i l i b r i a r e c r u i t m e n t c u r v e f o r p r e y s p e c i e s as d e s c r i b e d by Holling  (1973),  H a b e r ( 1 9 7 8 ) a n d o t h e r s t o show t h e r e l a t i o n s h i p b e t w e e n  h a r v e s t r a t e s and p o p u l a t i o n .  Maximum y i e l d s w i l l  o c c u r when r e c r u i t m e n t i s  36  F i g u r e 4.1:  Prey recruitment  c  curve  k/2  N*  Population in year = low s t a b l e  t  equilibrium  = unstable e q u i l i b r i u m (escape - max. N*  k k/2  harvestable  threshold)  surplus  = parent population density r e q u i r e d to produce = c a r r y i n g c a p a c i t y (upper s t a b l e e q u i l i b r i u m ) = population density with highest growth r a t e  instantaneous  = u n s t a b l e e q u i l i b r i u m below which n e g a t i v e recruitment leads to population e x t i n c t i o n  Y  M A X  37  maximum; t h i s o c c u r s a t a p o p u l a t i o n (w*)  somewhere between the c a r r y i n g  c a p a c i t y k and the p o p u l a t i o n a t which the f a s t e s t i n s t a n t a n e o u s dn/dt i s a c h i e v e d ( i . e . K/2). a p a r e n t s t o c k o f N*  individuals will  i n the f o l l o w i n g g e n e r a t i o n Consider  Removal  growth r a t e  o f the maximum h a r v e s t /max produced by Y  r e s u l t i n the same s i z e b r e e d i n g  ( i . e . N* ).  as  t+1  shown.  now the h a r v e s t y o f a p o p u l a t i o n p as shown i n F i g u r e  A t t h i s d e n s i t y , t h e r e i s a maximum h a r v e s t of i n d i v i d u a l s  (y*)  possible  t o the r e c r u i t m e n t w i t h o u t d i m i n i s h i n g the s i z e of the p a r e n t s t o c k . of l e s s  than the r e c r u i t m e n t ( i ^ ) w i l l  cruitment (y )  will  overharvesting  {Y>Y*) w i l l  d e p l e t e the subsequent  harvests  equal  Harvests  parent stock.  g r e a t e r than r e -  C l e a r l y , continued  d e p l e t e the p o p u l a t i o n t o t h e u n s t a b l e  brium a t c whereupon a l l s u r p l u s  4.2.  r e s u l t i n an i n c r e a s e i n the parent  p o p u l a t i o n the f o l l o w i n g g e n e r a t i o n and, c o n v e r s e l y , 2  stock  becomes n o n e x i s t e n t  equili-  (i.e. recruitment—  n a t u r a l m o r t a l i t y becomes the dominant v a r i a b l e , and the p o p u l a t i o n  0),  stabilizes  towards a low e q u i l i b r i u m d e n s i t y a t b. It ditional  i s c l e a r from such r e c r u i t m e n t curves  information besides  must know t o t a l  t h a t managers r e q u i r e a d -  a f i e l d d e t e r m i n a t i o n of r e c r u i t m e n t .  p o p u l a t i o n s i z e , a r e a l e x t e n t o f home r a n g e , the shape o f the  r e c r u i t m e n t c u r v e , and the v a l u e of k and o t h e r e q u i l i b r i a . cussed  i n more d e t a i l  accounted f o r .  A l s o , as  For w i l d l i f e managers u s i n g  o n l y r e c r u i t m e n t data t o  u n c e r t a i n t i e s which are not c o n s i d e r e d .  t h e r e f o r e , how e a s i l y o v e r h a r v e s t s The p r o c e s s  dis-  below, the e f f e c t s of s t o c h a s t i c change must a l s o be  a p o p u l a t i o n h a r v e s t these, a d d i t i o n a l c a t e g o r i e s of i n f o r m a t i o n substantial  They also-  might  represent  We can c o n c e p t u a l i z e ,  result.  i s more g r a p h i c a l l y v i s u a l i z e d i f p o p u l a t i o n  are c h a r t e d w i t h r e s p e c t t o t i m e .  establish  trends  F i g u r e 4 . 3 i l l u s t r a t e s an u n p e r t u r b e d ,  t o t a l l y d e n s i t y dependent, h y p o t h e t i c a l p o p u l a t i o n on a r e c o v e r y t r a j e c t o r y toward maximum d e n s i t y a t k.  The r e c r u i t m e n t component of each  seasonal  38  py  Py.  Population in year t+i  p  >  Population in year t F i g u r e 4.2: years.  E f f e c t s o f the h a r v e s t on r e c r u i t m e n t f o r s u b s e q u e n t p  parent population in year  y  harvest  Pyi  p a r e n t p o p u l a t i o n i n the y e a r f o l l o w i n g h a r v e s t YI p a r e n t p o p u l a t i o n i n t h e y e a r f o l l o w i n g h a r v e s t Y-  py*  Y*  2  p a r e n t p o p u l a t i o n i n t h e y e a r f o l l o w i n g no h a r v e s t ( h e r e where pop. d e n s i t y y i e l d s max. r e c r u i t m e n t ) - s p e c i a l ease h a r v e s t w h i c h e q u a l s max.  recruitment  39  7c  t  n  —7%  -  t+l  n  t+2 time in  t+3  —  t+4  years  F i g u r e 4.3: Annual and s e a s o n a l t r e n d s i n p o p u l a t i o n dynamics e x c l u d i n g random e n v i r o n m e n t a l e f f e c t s .  hunting season  b >  b-N* = g a i n from n a t a l i t y b-c - n e o n a t a l m o r t a l i t y e-N*= annual recruitment d-e - w i n t e r l o s s w*-e = o v e r h a r v e s t  c >| Population density  d >  V  XV**--..eern^bed.,^ate  M A N D F i g u r e 4.4: Annual c y c l e o f u n g u l a t e p o p u l a t i o n dynamics showing q u a l i t a t i v e e f f e c t s o f h a r v e s t i n g and e x c l u d i n g dynamic v a r i a b i l i t y ( m o d i f i e d a f t e r Dasmann, 1964, p.161). 0  40 cycle has been shaded to emphasize its nonlinear relationship with population density.  Recruitment is maximum at N * . Figure 4.4 is an enlargement of  one annual cycle portraying the effects of perturbation on the population trajectory.  Curve abcde represents the natural, unperturbed  stabilizing towards k as shown in Figure 4.3.  population curve  The increment e-N* represents  the number of individuals that could be removed from the population while maintaining the breeding population at N * . This "steady-state" trajectory is shown as curve a±>cde_j. Any overkill, through miscalculation of available yield or through inadequate control of hunting activity, will depress this curve further to e  2  and shift the population onto an extinction trajectory.  The need for corrective response is apparent.  Clearly, the level of harvest,  being the only variable in the yield equation over which managers have some immediate control, must be reduced in the following season to allow for recovery of the population to N * .  The longer the error remains uncorrected  the more d i f f i c u l t i t becomes to implement remedial action. In the foregoing paragraphs  I outlined the kinds of information  managers would require to determine harvest levels in a perfectly density dependent system-.  Such a system assumes, quite unrealistically, that " a l l  other things remain equal."  Random environmental  events, however, occur  commonly to affect natural populations in totally density independent ways. Of a l l global ecosystems, those in northern boreal and sub-arctic landscapes probably experience the greatest degree of environmental  change.  Extreme  seasonal cycles in these areas impose tremendous variability in the state of wildlife populations at any given moment, particularly the large mammal species.  In addition to climate and weather, floods, f i r e , accidents, and  habitat alteration by man can substantially influence the health and survivability of these species. What are the implications of stochastic change for •determining  41  potential harvests of wildlife?  Severe winters, for example, can take a  heavy toll of animals through starvation or increased predation.  Neo-natal  survival in the spring can be reduced through the malnutrition, disease or parasitism of mothers, increased exposure and predation of young, and so on. Such phenomena are common in wild ungulate populations. Unfortunately the frequency and intensity of the change is relatively unpredictable, at least in any simple quantitative sense.  Figure 4.5 illustrates the effects of  several kinds of environmental variability acting independently and in concert with one another.  Ungulate populations in the natural state undoubtedly  shift into periods of negative recruitment due to certain kinds of stochastic events.  Their survivability depends upon the inherent capability of the  species to recover in later seasons by various adaptive strategies to boost recruitment.  The amplitude of natural population variation due to such events  can be substantial.  Figure 3.4 showed the effect of random events on the  ungulate recruitment curve. Clearly, a preservationist harvesting policy must provide a buffer against stochastic events that negatively influence recruitment.  Under par-  ticularly adverse circumstances i t may be that, a particular population may experience a natural period of decline; in such cases there is no surplus available to the hunter.  This may indeed by the case for certain ungulate  species in British Columbia (Haber, 1978; Bergerud, 1978). From the theoretical evidence developed here and in Chapter III, i t becomes clear that s t i l l other kinds of ecosystemic variables may exist, which harvest policies do not incorporate nor respond to. These include natural or induced changes in carrying capacity of the habitat as well as the functional and numerical responses of interdependent species in the same or adjacent trophic levels (particularly predator-prey systems).  The con-  tinual removal of the "surplus" of one species could result in a competitor moving into the vacated food/space niche thereby creating a new carrying  42  P o p u l a t i o n curve under i d e a l  conditions  Figure 4.5: E f f e c t s o f random e n v i r o n m e n t a l events on annual u n g u l a t e p o p u l a t i o n c y c l e s i n n a t u r a l and hunted states: where... = reduced n a t a l i t y i n s p r i n g ; here equal t o annual r e c r u i t m e n t ;  b-b 1  h  1  to e  1  1  b  to e 1  b  3  to e 1  2  b. to e 1  4  = e f f e c t s o f reduced n a t a l i t y i n unperturbed system; = e f f e c t s o f reduced n a t a l i t y and i n c r e a s e d w i n t e r m o r t a l i t y i n unperturbed system; r:  e f f e c t s o f reduced n a t a l i t y and induced hunting m o r t a l i t y ;  = e f f e c t s of reduced n a t a l i t y , hunting and severe w i n t e r m o r t a l i t y .  capacity for the harvested population, possibly resulting in an accelerated decline as harvesting is perpetuated although no biological surplus remains. First and second order impacts such as this may affect other species and habitats in remote parts of the system (Talbot, 1975). One last but important point should be made here.  Harvest levels  that are calculated and applied as blanket policies rather than for distinct populations are ecologically unsound.  In species where definable units of  organization exist (i.e. a discrete population inhabiting a particular home range territory), harvestable surpluses must be determined for each.  That this  is critical is demonstrated by the following where harvesting policies were developed at a general, regional level and indiscriminately applied to many populations of the same species. The case of mountain goat is southern British Columbia is dramatic. Here, harvest levels have always been based solely on past harvest trends, a-t least up until 1978. The result has been an overall decrease in goat numbers as indicated by census data, field observations, and hunter harvest data:. In some cases local populations declined dramatically. These declines resulted primarily from overharvesting associated with increased and uncontrolled access (Phelps et al 1975, Foster 1976, also McRory pers comm.). "The overharvest was masked by a lack of detailed k i l l location information, and on a regional basis (i.e. the Kootenays) there was no downward trend in harvest success, sex ratios, or age class structure in the k i l l , ail potential indicators of overharvest.  An apparent decline in numbers of goats prompted a re-examination  of harvest information on a watershed basis which indicated an overharvest of some populations" (Petticrew 1979). Because wildlife managers had been monitoring harvest trend data on an aggregated basis at the regional level and, in addition, establishing harvest levels on the same basis;'' they were unable to detect the pattern of localized, sequential overharvesting of discrete goat populations as new  X 1964  —*  K  66  7*  *  68  r  7  ~K  70  "A™  ~~7*  72  *~  "  K  *  74  76  *  78  f i g u r e 4.6: Caribou P o p u l a t i o n d e c l i n e i n B r i t i s h Columbia by h e r d s ("after B e r g e r u d , 1978).  45 access was provided to unperturbed goat range.  The average provincial k i l l  from 1965 to 1973 was some 1,950 goats, dropping to 950 for the period 1974 to 1977. Bergerud (1978a) documented precisely the same pattern in his study of the provincial caribou decline. According to him, although overharvesting of several herds occurred in the 1960's, the overkill was not reflected in the total population until the next decade.  Hunting pressure on individual herds;  was apparently closely geared to the opening up of new caribou habitat to  j i  wheeled access.  This highly focused harvest sequentially depleted one herd ,  after another, and moved on to the next as each was overharvested and new access provided elsewhere (Figure 4.6).  Thus, even though distinct popula-  tions were being overexploited a high provincial total harvest was maintained until the 1970's when abruptly the reduction trend was manifested in harvest statistics. 4.3  Qualitative Impacts of the Harvest Up to this point I have been examining primarily the quantitative  impact of the harvest upon wildlife populations.  It is important to note,  however, that there are important qua!itative changes in the system that can be effected by increased mortality due to hunting.  Such changes are  not accounted for in conventional sustained yield calculations.  Whereas  the quantitative changes occur as direct reduction in population size, the qualitative ones occur as alterations, both direct and indirect, in the structure of the surviving population. Considering for a moment a theoretically ideal, single-species system i t is clear that sustained yield calculations do not incorporate relevant information on the age/sex distribution nor on induced effects on the social structure and behaviour of the exploited population. This latter effect is an extremely important aspect of the harvest  that is seldom addressed by wildlife managers.  Yet, there is good evidence  which suggests that the very act of the hunt itself has severe ramifications for the animal populations that survive.  Altmann (1956) for example, found  that at the opening of the hunting season elk undertook "evasive migration" (3 to 8 miles) which resulted in depressed rutting activity.  Similarly, Allen  et al.(1976) observed that elk moved from areas of low vegetation height to taller vegetation when vehicular access was created into virgin elk habitat. Lemke (1975) also monitored the effects of hunting on elk movement and found that migration-up to 6 miles occurred and use of heavy timber increased,. In a study of red- deer and chamois Douglas (1975) found that two years after the cessation of intensive hunting the animals became less wary and began to revert to preferred habitats.  Six years later some of the  hunting-induced effects were s t i l l observable in these populations.  In his  observations of red deer Geist (1971) found that habituation of hunted animals to secondary habitat causes a change in ecology and a resulting decline in population. He concluded that the worst and potentially fatal combination of harassment is hunting combined with hiking. sheep led Geist to conclude that  His later work on mountain  these animals readily alter their ecology  in response to experience and, where protected from hunting,.will exploit manmade habitats and accept people.  Sheep do not disperse readily and, when  hunted, they are easily pushed to extinction through confinement to safer but sub-optimal habitats (Geist, 1975a). In testimony before the Mackenzie Valley Pipeline Inquiry, Geist categorized the effects of disturbance, both through hunting and other forms, into three types:  (1) increase in required energy budget of the individual;  (2) illness, death, or reduced reproduction; and (3) avoidance of preferred habitat (Geist, 1975b). The qualitative nature of the harvest, i.e. the "style" of the  hunt, also appears to bear an important relationship to the social structure and well-being of the exploited population.  In "sport" or recreational  hunting, as a prominent example, the harvest is sometimes intentionally concentrated on particular animals in the population - the largest, prime males of the species.  This is the so-called "trophy hunting", which can be charac-  terized as a non-random, highly selective removal of a particular age/sex cohort of the population.  There is some evidence to suggest that the removal  of the large, mature males in a big game population produces alterations that can range from seemingly t r i v i a l effects to serious negative changes.  In  bighorn sheep, for example, McCann (1956) detected increased wariness in rams due to hunting activity.  Similarly, De Forge (1976) observed that bighorn  sheep lack the capability to colonize new range and that human disturbance leads to range abandonment,and overcrowding on unperturbed parts of the range which, in turn, resulted in increased stress and abnormal social relations. Geist (1971) also has speculated that disturbances caused by hunting may set up avoidance behaviour in sheep so that portions of traditional range become eliminated from collective herd knowledge. The maintenance of home range tradition in the older members of both sexes has particular importance here.  The knowledge of key winter and  summer ranges, calving grounds, escape terrain, and other critical habitat appears to be stored and implemented by the oldest animals in the population (Geist, 1971).  The failure to establish crucial patterns of migration has  been documented in caribou reintroduced t o suitable habitat but lacking in home range knowledge (Bergerud, 1971).  This concept has particular relevance  for the management of those big game species wherein the hunt is focused on the older, "trophy" males.  The loss of tradition in either sex, especially  in those species where the seasonal ranges o f males and females can differ, could have serious deleterious effects on the population (Cowan, 1974).  Cowan a l s o s u g g e s t s  that the l o s s of experienced  leaders during the  random e x p l o i t a t i o n o f animal  p o p u l a t i o n s can d i s r u p t r e p r o d u c t i v e  In s p e c i e s which  synchronous  have adopted  ary mechanism to minimize of breeding  behaviour.  r u t t i n g and c a l v i n g as an e v o l u t i o n -  p r e d a t i o n on y o u n g ( B e r g e r u d ,  t r a d i t i o n such as a s h i f t t o a s y n c h r o n o u s  r e s u l t i n increased neonatal m o r t a l i t y through to  non-  1974)  mating  prolonged  the d i s r u p t i o n and c a l v i n g c o u l d  exposure  of young  predators. G e i s t has a l s o s u g g e s t e d  t h e b r e e d i n g s t o c k may over the long term,  t h a t the harvest of trophy animals  from  r e s u l t i n d e p l e t i o n o f the p o p u l a t i o n gene pool  cumulative  genetic impoverishment  of the herd.  and,  Moreover,  Haber (1977) has i n d i c a t e d t h e p o s s i b i l i t y t h a t h a r v e s t i n g o f t r o p h y s p e c i m e n s p a s t p r i m e b r e e d i n g a g e may dation also utilizing wolves  s h i f t a compensatory  those older animals  i n t o a noncompensatory form  pre-  where  a r e f o r c e d to prey upon b r e e d i n g age i n d i v i d u a l s . Another  impact  o f t r o p h y h u n t i n g h a s s h o w n up r e c e n t l y i n s o m e o f  the p r o v i n c e ' s r e s i d e n t moose p o p u l a t i o n s . ditional imbalance  hunting r e g u l a t i o n s which  emphasized  C h i l d ( 1 9 8 0 ) has n o t e d t h a t  tra-  t r o p h y h u n t i n g has c r e a t e d  an  between the sexes, g e n e r a t i n g a p o p u l a t i o n s t r e s s t h a t , i n t u r n ,  r e s u l t e d i n lower p r o d u c t i v i t y (more d r y cows), mortality of late-born calves. mature, experienced duced  component of wolf  b u l l s from  recruitment because  asynchronous  r u t , and  In o t h e r w o r d s , t h e c o n c e n t r a t e d  higher  removal  of  t h e b r e e d i n g p o p u l a t i o n has r e s u l t e d i n r e -  o f t h e low s u c c e s s r a t e o f i n e x p e r i e n c e d young  bulls. In summary, and w h i l e n o t c o n c l u s i v e , t h e t h e o r e t i c a l and e m p i r i c a l evidence  suggests  t h a t t h e f o l l o w i n g p h e n o m e n a may  r e s u l t from hunting  in  general: 1.  Overall population decline;  2.  Abandonment Of r a n g e ; optimal range;  c o l o n i z a t i o n and o v e r c r o w d i n g  o f sub- *  3.  Increased  s t r e s s , w a r i n e s s , and abnormal s o c i a l  behaviour;  4.  S h i f t o f p o p u l a t i o n i n t o a l o w - s t a b i l i t y domain where p r e d a t o r s become l i m i t i n g f a c t o r ;  and from trophy h u n t i n g i n p a r t i c u l a r : 5.  D i s r u p t i o n o f r e p r o d u c t i v e b e h a v i o u r due t o l o s s of e x p e r i e n c e d 1eaders;  6.  Loss o f home range  7.  Long-term g e n e t i c impoverishment o f the p o p u l a t i o n ;  8.  I n c r e a s e d p r e d a t i o n upon b r e e d i n g - a g e i n d i v i d u a l s as c o m p e t i t i o n f o r o l d e r , prime specimens i n c r e a s e s .  tradition;  Where n o n - s e l e c t i v e h u n t i n g a l o n e c o u l d p o t e n t i a l l y produce t h e s e e f f e c t s i n the hunted p o p u l a t i o n , - t h e l a s t f o u r l i s t e d are l i k e l y to be more e a s i l y generated through a h a r v e s t i n g regime which f o c u s e s on mature male specimens.  One cannot prove c o n c l u s i v e l y w i t h the e v i d e n c e t o date t h a t a l l  these e f f e c t s w i l l  happen t o a l l hunted p o p u l a t i o n s ; nor i s such a c o n t e n t i o n  a p p r o p r i a t e to t h i s a n a l y s i s .  The p o i n t i s t h a t some of these e f f e c t s have  been observed to o c c u r i n one s i t u a t i o n or a n o t h e r , a l l seem i n t u i t i v e l y r e a s o n a b l e , and a l l c o r r e l a t e s t r o n g l y t o developed t h e o r y which t e l l s what we might e x p e c t .  us  T h i s may o r may not have p a r t i c u l a r r e l e v a n c e f o r  h a r v e s t i n g regimes which a r e drawn up f o r the p r o v i n c e as a whole and i n which the g e n e r a l p o l i c y i s to e x p l o i t the w i l d l i f e r e s o u r c e u n t i l are encountered.  problems  I t does, however, have immediate r e l e v a n c e to a s i t u a t i o n  wherein hunting a c t i v i t y must take p l a c e i n accordance w i t h a p o l i c y o f p r e s e r v i n g n a t u r a l communities or n a t u r a l  systems.  Before d e v e l o p i n g a s e t o f management g u i d e l i n e s which d e r i v e from those p r i n c i p l e s , i t i s n e c e s s a r y sizi  to d e s c r i b e the w i l d l i f e community i n  Spat-  Park to ensure t h a t i t s p r o p e r t i e s are c o n s i s t e n t w i t h those o f the t h e o -  retical chapter.  and o t h e r n a t u r a l systems d e s c r i b e d above; t h i s  I do i n the f o l l o w i n g  CHAPTER  V  Spatsizi - A Description of the Wildlife Component  5.1  Spatsizi Wildlife Communities - An Overview In this chapter I intend to describe briefly the relevant charac-  teristics of the Spatsizi wildlife community and demonstrate that the properties and principles discussed in preceding chapters have applicability here. The Spatsizi territory, and by this I mean the land in and adjacent to Spatsizi Plateau Wilderness Park, is one of chose regions in the northern part of the continent where large tracts of wilderness and their indigenous wildlife populations have been preserved relatively intact.  The wildlife  systems of these boreal forest and alpine zones are characterized by the f o l lowing features: sers in winter;  (I) abundant small mammals;  (2) many -gra.ze.rs become brow-  (3) 80% of the nesting bird population migrates; and (4)  the complete food chain is present including large carnivores (Siderits Radtke, 1977).  This last, the presence of naturally functioning large  carnivore predator-prey systems, is probably the salient feature of Spatsizi Park. The most visually prominent lifeforms in this region are the wild ungulates (moose, caribou, mountain goat, thinhorn sheep, and mule deer), their important predators (wolf, grizzly bear), and several other large carnivores and omnivores (black bear, lynx, wolverine, coyote, fox and so forth).  Not only is this assemblage diverse but, relative to other areas of  British Columbia, overall populations of these species are generally high. The trophic web (Figure 5.1) demonstrates the interrelationship among northern boreal species and how the large mammals are functionally integrated into a much larger community of smaller mammals, birds^ and vegetation^. While seemingly complex, i t is only a simplistic representation of the i n t r i cacies of such a natural system.  Each species component, for example, can be  further disaggregated into an entire "sub-assembly" with many interacting variables.  Figure 5.2 depicts the case for moose (Alces  aices).  Clearly, an  infinite amount of information would be required for the perfect understanding of the dynamics of any one species, let alone an entire ecosystem. Because the Spatsizi is a land which is subject to the extreme vagaries of sub-arctic weather, i t follows that the dynamic variability in the distribution and abundance of resident wildlife populations is high. Populations of ungulates move from winter range to summer range with migrations occurring both vertically and laterally in space.  Aggregations of  animals occur in the f a l l for mating; dispersion occurs for calving in the * Osmond-Jones et al (1977) l i s t 125 species as observed and confirmed, 11 as hypothetical, and 12 observed by other naturalists. O L  h  Pojar (1976) recorded 370 species of vascular plants in the Gladys Lake Ecoreserve in the center of the Park.  52  N  )  Mouse  I'i,|in l i i l . i o n  M u r l a l i Ly  / N  Size  Rate  /  N a t a l it y K a u  ~7JT  Zb_ Km i g r a t i or Immigration  Wi n t c r M o r t a 1 i Ly  l l u t i t i ng  0  5  Sociai O r g a n i z a t it.  —^  >1  A  Prcgnaney Rate  "7fT  Winter-  r)  ~  1WI1 Natali ty  As ,  Y e a r l i ng  j^arusi t e s  ~7fC  Calf Mortali ty  Pregnancy Rate  Kawn  Decs i ti  £1  Ovu1 a t i o n  0 is t r i b u ti on  Rate  Y e a r l i ng  I  I  N a t a l i l.y  j  j  Aduit Natal i ty  Z2T All-.i! t Pregnancy, Hate  Y e a r 1 I tig Ova I a t i t m Hate .  Adu I t O v u I at. i o n .Rate  POPULATION DFSPOftSKS HAM]TAT Avai1abIc  RESrONSES  Quality Annua i  Forage  of  Knergy  Forage  ~7f\  W a i J a b i Ii t y  Furagc  Plant  Condit ion  'Juani.i t y u f rorafe  ~7SL C o m p e l . ; i; i v e Grazing  Annua i { l o n d i *, i ont:  5.2  Population and habitat variables affecting density of m o o s e {Alces a Ices ). Adapted from Gross, 1972.  spring.  To some extent their natural predators move with them, presumably  attracted by high densities or susceptible individuals as the season dictates. Major shifts in the distribution and seasonal ranges of some species have been observed (Haber, 1978). Of a l l the resident species in the park those comprising big game have drawn the most attention from hunters and, more recently, the general public with their concerns related to trophy hunting in Spatsizi.  Caribou,  sheep, goat and grizzly are the star attractions for hunters with moose, perhaps, f u l f i l l i n g a secondary role.  The competition between hunters and pre-  dators for ungulates over the years has on occasion been substantial, peaking during the 1950's with aerial poisoning of wolves.  What information exists  to more clearly define the predator-prey relationships that characterize the Spatsizi wildlife community? According to Haber (1978) there are three or four wolf packs which live in the upper Stikine-Spatsizi area.  The principal food of wolves in  northern regions is moose, even when other prey are numerous (Rausch, 1969). This is in accord with Bergerud's (1978) and Haber's (1978) findings in the Spatsizi which demonstrate that moose is the main food source with caribou, sheep, goat, mule deer, and beaver making up the remainder.  Their information,  however, suggests that some seasonal preference for caribou exists during the summer and, particularly, during calving season when caribou young are susceptible to predation.  In addition, as noted earlier, the functional response  of wolves is dependent upon prey density and, therefore, under certain conditions prey capture will be related tc prey availability and vulnerability. Caribou, on the other.hand, are food not only for wolves but also for grizzly and, to a lesser extent, wolverine.  The heaviest mortality due  to predation for these animals occurs during spring calving when grizzlies become a significant factor in calf mortality. Bergerud s data (1979) seems !  55  to indicate a mortality rate in the f i r s t year of 90% for Spatsizi caribou calves compared to 80% for other northern populations.  He suggests that  grizzlies are as important a mortality agent as wolves for caribou although this has not been empirically demonstrated.  Field observation, however,  suggests that grizzlies form a search image for caribou during the calving period in late spring and can be successful predators of new-born caribou. The observation here is that predator-prey relationships between the larger mammals of the Spatsizi are characterized by two dominant predators utilizing moose as principal prey but also caribou as preferred alternate food at particular times of the year.  The resulting system dynamics are  likely complex, and highlight an important issue for the management of game harvests. In relating features of Spatsizi wildlife resources back to the important characteristics of ecological systems described in Chapter II, i t is clear that dynamic variability and uncertainty are high, and that the system which comprises the big game populations in the area is ordered - that i s , a number of species co-exist which are functionally interconnected so that the population response and behaviour of one can affect any one or more of the others.  The one additional property not discussed to this point is the rele-  vance of the multiple equilibria predator-prey model to Spatsizi wildlife. Is. there evidence to suggest its applicability in this particular area? Research carried out in the Spatsizi by Haber (1978) on the wolfungulate system and by Bergerud (1978) on caribou seems to support the concept of a non-linear relationship between prey and predator densities and the existence of high and low stability domains for caribou.  Interpretation of  some aspects of the model and field information differs between these two biologists but there seems to be .general agreement on the principles involved. Haber's extensive work on arctic caribou led him to certain, conclusions about distribution and abundance of populations that may have  relevance to the woodland caribou of northern British Columbia.  According  to him, caribou occupy discrete home ranges centered around key feeding and breeding grounds and extending into less important but s t i l l accessible rangeland.  Piecing together the historical record led him to believe that  population levels of individual herds rise and f a l l in a cycle of approximately 60-90 years.  The availability of food in the range appears to be the  controlling factor of population highs with predation as a secondary factor. When food competition becomes too intense dispersal from the expanding herd results in the migration of some individuals into neighbouring ranges where, ideally, the resident populations are low enough to accommodate the immigrants. Haber's data suggests adjacent caribou herds as being out of phase with one another with respect to population levels.  Such an arrangement could  provide for the dispersal from overcrowded ranges into underutilized ones so that large-scale starvation does.not occur.  Haber suggests that i f dispersal  does not occur then a severe food crash results in collapse of the range and mass starvation.  This is in disagreement with Bergerud (1979) who argues t-hat  food is seldom a limiting factor and that caribou populations generally are kept below high equilibrium densities bv poor recruitment and high natural mortality. Enough field investigation has not been carried out to show that caribou populations in the Spatsizi behave in the same way.  Nevertheless, we  do know that several, well-defined rutting grounds exist in the area which suggests that the conventional approach of defining caribou herds by their ruttings grounds may be appropriate.  Bergerud and Haber have concluded that  Spatsizi caribou populations can be broken into two or three distinct herds. Recent radio-collar tagging by Hatler (1982 pers comm.), however, indicates that individual animals seem to have a much greater freedom of association than hitherto supposed and do not necessarily re-unite with the same breeding  population as they did the year before.  Hatler's sample is very small (15  animals originally tagged, 10 surviving as of this writing) but he is beginning to suspect that the Spatsizi caribou belong to one large amorphous population that fragments and aggregates in clusters somewhat randomly on a seasonal basis.  More work needs to be done to prove or disprove this contention.  Certainly, dispersal of some herds has been recorded and seasonal migration of some of them appears to be substantial.  Major and apparently permanent  shifts in distribution have also been recorded in recent years by guide outfitters in the area (Haber 1978) although these displacements may be related more to railroad construction and increased hunting pressure than natural causes. Haber (1978), drawing from historical caribou data and simulation modelling, goes on to suggest that i t is likely that woodland caribou recruitment curves exhibit the same topological features as other predator-prey systems that have been investigated (Haber, 1977; Walters et a l , 1978; Holling et al , 1976; Peterman and Gatto, 1977; Brand et a l , 1976).  While not docu-  mented in the case of the Spatsizi, i t nevertheless suggests possibilities that must be considered by managers i f the policy mandate for the park is to be upheld.  Figure 5.3 depicts Haber s recruitment curve for arctic caribou. !  Its purpose here is to underscore the possibility that caribou population dynamics are characterized by a high and low density stability domain.  On the  basis of studies in Denali National Park, Haber (1978) argues also that a similar recruitment curve topology potentially exists for moose in the Spatsizi. Haber's interpretations about such curves echo the ones made here in earlier chapters.  As herd density approaches either equilibria, the net  annual- increments to the population decrease, becoming negative below the lower equilibrium and above the upper. In the case of arctic barrenlands caribou (Figure 5.3), there is a high stable equilibrium at densities around 5 animals per square mile  58  intraspecific competition 1  Recruitment low  unstable (escape threshold)  high stable:  •k population density in caribou per sq. mi,  Figure 5.3: Density dependent recruitment curve for caribou. Shaded area represents 'predator pit' where depensatory effects of wolves will maintain prey densities @ low stable equilibrium. Heavy dotted 1ine represents Bergerud's model where predation leads to extinction at *. Values for c and k have beenjderived by Haber (1977) from historical highs and lows of caribou fluctuations in the Arctic. See also Bergerud (1979).  59 caused by increased intraspecific competition for food and reproduction and, hence, lowered fecundity levels in the population.  Skoog (1968) believes  that recruitment rates decrease at high densities due to more erratic herd movements, more frequent separation of young and mothers during large migraitions, and deteriorating range conditions. Haber suggests, too, that rapidly depleting food supply at high densities triggers dispersal and resident populations drop.  In this situation, herd densities would tend  to fluctuate at levels between k and c (Fig.5.3) at periodicities geared to range buildup and deterioration.  Populations driven down to densities  just above the escape threshold at c (one animal per square mile) will tend to rebound to the high equilibrium level. Once declining populations reach the unstable equilibrium at one individual per square mile then they are set on a new trajectory caused by negative recruitment and will stabilize at some new low equilibrium b (Haber, 1977).  Bergerud (1979) suggests, on the other hand, that a narrow stability  domain exists at c and that predators can stabilize herd densities at this population.  Increased mortality at this level, however, may result in a nega-  tive trajectory culminating in extinction (asterisk in Figure 5.3).  Haber's  argument against the extinction model is that virtually a l l the remnant caribou herds in Alaska, Yukon and Northwest Territories have continued to persist at low densities in presence of wolves. Haber believes that natural predation alone cannot be a causative agent in caribou declines from high density levels - some other factor such as overharvesting is necessary to reduce herd levels to escape threshold at c. Once below this level, however, predation can be a primary factor in maintaining low densities (0.5 - 1.0 caribou/mi. ) until such times as immigration from a neighbouring herd boosts the density level above the escape threshold and into the high equilibrium stability domain.  A beneficial result of this  "depensatory" effect of predators upon ungulates is that time is provided for  60 the range condition to recover. Haber is careful to note that sole reliance on recruitment data for setting harvest levels without any determination  of population density  could lead to the conclusion that population levels were low rather than high.  "North American wildlife biologists have long argued that ungulate  populations can be managed solely on the basis of sex-age trend i.e. without any information on population size (density), but numerous recent major ungulate declines throughout the north and the analyses in this report as well as Haber et al (1976) and Haber (1977) indicate otherwise" (Haber 1978, p.64). Haber (1978) also warns against the possibility that "sampling errors alone could mask the smaller changes and hence indication of further decline in numbers once a population is dangerously close to its lower equilibrium." Earlier in this chapter I have attempted to show that the wilderness communities of the Spatsizi share the major features of most ecological systems; high dynamic variability and uncertainty, and ecological order.  In  the latter part I have presented some evidence to argue that, while not conclusively demonstrated, i t is likely that the existing wolf-ungulate system is characterized by the non-linearity and multiple equilibria which are found in other predator-prey  systems.  This being the case, i t can then be argued  that the key principles which have been identified in Chapter II as essential features of naturally functioning ecosystems apply as well to the Spatsizi area.  It remains now to formulate a set of guidelines for the park which em-  body the identified ecological principles and the main features of the resident wildlife community in a cohesive management framework consistent with the Order-in-Counci1.  CHAPTER  VI  Ecological Guidelines for Spatsizi Wildlife Management  6.1  Guidelines for Wildlife Management As noted earlier wildlife populations are affected by a host of  driving variables that results in dramatic fluctuation in numbers and spatial distribution.  Intraspecific and interspecific competition (including pre-  dation) can influence these changes in a non-linear, density-dependent manner. The theoretical evidence suggests that multiple stability domains and extinction thresholds characterize wild ungulate population dynamics;. These features have relevance in management schemes which permit the harvest of wildl i f e and yet are intended to minimize impacts upon the remaining populations. What should be clear from the preceding chapters is that i f wildl i f e management for Spatsizi Park is to achieve the goals laid out by the Order-in-Counci1  then i t must adopt a 'systems approach', incorporating•both  the operative principles of natural ecological systems as well as the main features of the Spatsizi wildlife community.  62 T h e s e p r i n c i p l e s may be summarized as f o l l o w s : 1.  The n o n - l i n e a r r e l a t i o n s h i p s and m u l t i p l e e q u i l i b r i a which c h a r a c t e r i z e p r e d a t o r - p r e y systems must be a c c o u n t e d f o r ;  2.  Dynamic v a r i a b i l i t y must be a c c o u n t e d f o r i n e s t a b l i s h i n g harvest levels;  3.  The i n h e r e n t o r d e r o f w i l d l i f e communities  4.  U n c e r t a i n t i e s must be r e d u c e d o r overcome by a p p l y i n g approp r i a t e safe-guard measures.  must remain  intact;  A t t h e o u t s e t , i t s h o u l d be c l e a r t h a t t h e most a d m i n i s t r a t i v e l y s i m p l e , l e a s t e x p e n s i v e , and p r o b a b l y most e f f e c t i v e way o f p r e s e r v i n g t h e : S p a t s i z i w i l d l i f e base i n t a c t and p e r m i t t i n g t h e n a t u r a l s u c c e s s i o n o f events w i t h i n t h e park i s t o p r o h i b i t a l l human u s e . A t o t a l ban on human v i s i t a t i o n would a c h i e v e t h e p r e s e r v a t i o n i s t i n t e n t o f t h e O r d e r - i n - C o u n c i l . The p r o v i s i o n f o r l i m i t e d h u n t i n g and f i s h i n g r e p r e s e n t s , t h e r e f o r e , a dilemma. To what e x t e n t can t h e s e a c t i v i t i e s o c c u r b e f o r e t h e i n t e g r i t y o f t h e i n d i genous communities  and n a t u r a l s u c c e s s i o n i s i m p a i r e d ?  T h i s b r i n g s us i m m e d i a t e l y t o t h e q u e s t i o n o f f o r m u l a t i n g b i o l o g i c a l ly 'safe' harvest l e v e l s .  What i n f o r m a t i o n i s r e q u i r e d by managers t o ensure  t h a t h u n t i n g does n o t d i s r u p t t h e i n t e g r i t y o f t h e S p a t s i z i w i l d e r n e s s ecosystem?  The maximum a l l o w a b l e h a r v e s t o f any s p e c i e s c l e a r l y cannot  exceed  t h a t p a r t o f t h e n e t annual i n c r e m e n t as c a l c u l a t e d a f t e r t h e e f f e c t s o f n a t u r a l m o r t a l i t y and i l l e g a l h u n t i n g have been a c c o u n t e d f o r . A d d i t i o n a l l y , the 'take' has t o be low enough so t h a t p r e d a t o r r e s p o n s e p a t t e r n s a r e n o t s i g n i f i c a n t l y a l t e r e d and t h e b e h a v i o u r o f p o p u l a t i o n s remains  unperturbed  by h u n t i n g a c t i v i t y . The a c c u m u l a t i o n o f trie d a t a n e c e s s a r y t o meet t h e s e o b v i o u s l y r e p r e s e n t s a s u b s t a n t i a l t a s k f o r w i l d l i f e managers.  requirements Nevertheless,  i t forms t h e f i r s t e s s e n t i a l g u i d e l i n e : F o r .each hunted s p e c i e s t h e f o l l o w i n g i n f o r m a t i o n must be a c q u i r e d ;  t r e n d s and p r e s e n t s t a t e o f p o p u l a t i o n s i z e  and age/sex d i s t r i b u t i o n ; n e t annual i n c r e m e n t ; ext°nt o f range; d e t e r m i n a t i o n  63  of c a r r y i n g c a p a c i t y ; and behavioural hunting  responses  of surviving populations to  harassment and l o s s o f herd e x p e r i e n t i a l knowledge.  I t i s important  t h a t t h e f i r s t two c a t e g o r i e s o f i n f o r m a t i o n be a c q u i r e d s y s t e m a t i c a l l y and annually; the other categories represent  i n f o r m a t i o n o f a more general  that i s n o t n e c e s s a r i l y s u b j e c t to annual t i o n such  as t h a t p e r t a i n i n g t o range  variables which a f f e c t vegetation  variation.  Nonetheless,  nature  informa-  c o n d i t i o n and i n f l u e n c i n g random  (food o r c o v e r ) must be r e g u l a r l y o b t a i n e d  so t h a t p r e d i c t i o n s o f s t o c h a s t i c c h a n g e i n t h e h a b i t a t c a n be made. The  e f f e c t s o f other v a r i a b l e s such as f e c u n d i t y o f t h e p o p u l a t i o n ,  natural m o r t a l i t y due t o s t o c h a s t i c events,  predation, and i l l e g a l  are a u t o m a t i c a l l y i n c o r p o r a t e d i n t h e p o p u l a t i o n census recruitment, but only i f this information i s developed (annually f o r those species which reproduce range  and determination o f f o r each  once a y e a r ) .  c a r r y i n g c a p a c i t y i n t h e above i s important  generation  The i n c l u s i o n o f  as i t i s e s s e n t i a l that  the upper l i m i t s t o p o p u l a t i o n d e n s i t y be a s c e r t a i n e d even i f o n l y mately. nual  T h i s i n f o r m a t i o n then enables  increments  sities  managers t o determine  approxi-  i f low n e t an-  r e p r e s e n t high p o p u l a t i o n d e n s i t i e s o r low population  ( i . e . d e n s i t i e s near t h e escape  librium).  hunting  den-  t h r e s h o l d o r below a low s t a b l e e q u i -  In other words, the determination  of density alone  i s inadequate;  knowledge o f the upper l i m i t s t o population s t a b i l i t y w i l l b e t t e r a i di n establishing the overall state of the population. An a n a l y s i s o f t h e a b o v e i n f o r m a t i o n w o u l d p r o v i d e t h e b a s i s f o r c a l c u l a t i o n o f t h e maximum a l l o w a b l e h a r v e s t ; t h a t i s , t h e number o f i n d i v i d u a l s t h a t m i g h t be removed w i t h o u t the p o p u l a t i o n . Another  a c t u a l l y d i m i n i s h i n g the growth r a t e o f  I t i s , a t this stage, primarily a quantitative  determination.  q u a l i t a t i v e g u i d e l i n e i s necessary' t o s t r u c t u r e t h e harvest so that  w i l d l i f e communities a r e t o remain  u n a l t e r e d ; h a r v e s t i n g must be u n i f o r m l y  r e g u l a t e d so t h a t n a t u r a l l y o c c u r r i n g age/sex d i s t r i b u t i o n s o f each are preserved.  This requirement  would n e c e s s a r i l y l i m i t trophy  population  hunting  64  to the extent consistent with the naturally occurring age/sex distribution. Part of the total harvest, therefore, must consist of younger males and an appropriate distribution of females. This leads to consideration of the effects of dynamic variability in determining harvests.  The differences in social behaviour between each  species have implications for the way harvest levels are established. Moose, for example, are usually spaced more homogeneously throughout their range and typically do not travel more than a few miles in any one year.  Caribou  tend to aggregate in high-density clusters for certain parts of the year and to migrate or disperse dozens i f not hundreds of miles at other times.  Sheep  and goat, on the other hand, aggregate more or less continuously in small numbers and wander over home ranges of a much more limited extent than caribou. Thus, the variation in movement and population size over the range and throughout the year suggests' that harvest strategies should be geared differently to each species.  In some cases discrete identifiable populations exist, in others  populations are so mobile or so widely dispersed that no coherent 'unit' can be delineated.  This leads to the derivation of a third guideline for the harvests  of Spatsizi wildlife:  where applicable, harvest levels for each species must  be established for discrete communities or populations, and not on a provincial, regional, or otherwise arbitrary basis. The dynamic variability inherent in the Spatsizi wilderness ecosystem clearly necessitates a flexible approach to the establishment of annual harvest levels.  Assuming even .relatively stable population densities, random  environmental  change can act so as to unpredictably increase or diminish net  annual increments; the most dramatic example being in-migration or dispersal. In some years, for example, when populations are near optimum densities, recruitment is high, and the preceding winter has been a relatively benign one, hunters may be able to enjoy high harvests (assuming, of course, that such  harvest levels are otherwise properly determined).  In other years, due to  any number of extenuating circumstances hunters may have to accept minimal 'takes' or may even have to forego harvesting opportunities altogether until a subsequent season.  Harvest levels cannot, therefore, be set on some arbi-  trarily conceived fixed level (say, a certain percentage of the population). This leads logically to the derivation of a fourth guideline for management: fixed yield policies (i.e. harvest levels established for more than season) must be eliminated and pulse harvesting policies instituted. With respect to the management of predators in the park, the Orderin-Council makes i t clear that natural processes are to remain unaltered. Controlling predators in order to boost ungulate populations for increased hunting would necessarily alter the balance between predators and ungulates, thereby violating the policy requirement.  My f i f t h guideline i s , therefore:  predator control should not be practised. This immediately begs the question of what might constitute appropriate restorative strategies in the light of declining ungulate populations. If the effects of human use of Spatsizi wildlife resources have already created a situation wherein natural predation is depressing ungulate recruitment, then is i t not appropriate to intercede once more in an attempt to correct the imbalance by controlling predators?  It may be, but the decision must be based  on a very clear understanding of the dynamics of the system and the efficacy of predator reduction to achieve the desired results. An important requirement would be that managers determine; the causal relationship between ungulate declines, predation, and human overkill.  His-  torical population trend information is vital to ensure that a true decline is underway for, as Haber (1978) suggests, certain low-density populations may have always existed below the escape threshold as the initial colonization of a particular range may have been carried out in the presence of natural predators which continuously maintained the herd at low density.  Such a  66 s i t u a t i o n ; w o u l d demonstrate the n a t u r a l , l o n g - t e r m v i a b i l i t y o f p r e d a t o r - p r e y systems (assuming o t h e r v a r i a b l e s  low-density  remain r e l a t i v e l y c o n s t a n t )  thus o b v i a t i n g any immediate cause f o r c o n c e r n . A n o t h e r c a t e g o r y o f i n f o r m a t i o n t h a t must be developed i n t h i s s i t u a t i o n i s the r e l a t i v e d e n s i t i e s of the p e r t i n e n t p r e d a t o r ( s ) a l t e r n a t e prey.  In the S p a t s i z i  wolves to a i d i n b o o s t i n g  the c a l l  has been made f o r a r e d u c t i o n  caribou recruitment.  however, the f u n c t i o n a l response  of wolves  Moose i n the S p a t s i z i  in  As I have argued e a r l i e r ,  towards c a r i b o u w i l l  upon the abundance o f the l a t t e r r e l a t i v e t o the p r e d a t o r ' s i n t h i s c a s e , moose.  and t h e i r  be dependent  preferred  food(s),  a n d , more g e n e r a l l y , i n o t h e r  p a r t s o f n o r t h e r n B r i t i s h Columbia are a p p a r e n t l y d e c l i n i n g i n numbers to o v e r h a r v e s t i n g  and low r e c r u i t m e n t ( B e r g e r u d ,  b i l i t y of t h i s food source may be r e s p o n s i b l e ,  1978).  due  The d i m i n i s h e d a v a i l a -  i n p a r t , f o r the  depensatory  e f f e c t s on c a r i b o u i f wolves a r e being f o r c e d t o s h i f t t h e i r p r e d a t i o n t o the latter.  Thus, h u n t i n g p r e s s u r e on both these main f o o d s p e c i e s  c o u l d be a c -  c e l e r a t i n g the impacts o f p r e d a t i o n upon b o t h . I have d e s c r i b e d t h e above p o s s i b i l i t y a t the r i s k of g r e a t l y plifying reality.  The e s s e n t i a l  p o i n t i s t h a t the dynamics  sim-  o f the p e r t u r b e d  system may be r e l a t i v e l y complex and, t h u s , the i n f o r m a t i o n needs o f managers t o make a sound d e c i s i o n w i l l  be s u b s t a n t i a l .  In f a c t , not o n l y i s the amount  o f data s u b s t a n t i a l , but the d i f f i c u l t y o f o b t a i n i n g t h i s d a t a i n a r e l i a b l e form i s  immense.  To assemble  it will  not be an easy t a s k ; y e t i f h u n t i n g  is  t o be a l e g i t i m a t e use o f the park then e x t r a o r d i n a r y management e f f o r t w i l l be n e c e s s a r y  t o ensure t h a t park o b j e c t i v e s are not c o n t r a v e n e d .  A final  n o t e ; i f d e c l i n e s are r e a l and the r e s u l t o f human impact  then i n no way must p r e d a t o r c o n t r o l be implemented w i t h o u t a p r i o r s u s p e n s i o n of the a c t i v i t y t h a t p e r p e t r a t e d the d e c l i n e s i n the f i r s t i n s t a n c e . i n o t h e r words, must cease and the e f f e c t s m o n i t o r e d long b e f o r e any to reduce p r e d a t o r s  i s made.  Hunting, decision  The preceding rules comprise what I believe to be a framework for establishing a level and structure for game harvests that would be consistent with  the Order-in-Counci1.  An obvious implication arises here; i f the neces-  sary information as described above is not available to park managers, for whatever reasons, then there is a serious risk that continued hunting will violate the "preservation" objective of the Order-in-Council.  Because the  uncertainties in this situation are so great, the cessation of hunting appears to be the only viable policy that will ensure that hunting does not impact natural communities. Assuming, on the other hand, that a substantial effort is mounted to collect reliable biological information there s t i l l remains the question of how to cope with uncertainties inherent in the collection and interpretation of the data.  Methods used to inventory populations, for example, might introduce  significant error into the results (say, through double-counting); home range territories may be incorrectly defined; the effects of dispersal or immigration may be wrongly determined; the psychological and social effects of hunting harassment on wild animals s t i l l improperly understood - these and many more unpredictable sources of error obligate managers to incorporate, conservative measures in developing harvest strategies for the park. The starting point seems an obvious one.  Harvest levels that are  suggested as permissible by careful analysis of the relevant data s t i l l do not eliminate a certain degree of risk.  Reliability of the data gathered may be  insufficient to guarantee that continued hunting will have no significant effects on game populations.  A further reduction of harvest levels-, at this  point, serves to decrease the risk involved. Much of the uncertainty arises because the impacts of the harvest on wildlife populations are, for the most part, not well understood. An obvious approach for management would be the careful and rigourous monitoring  68  of two separate big game communities; one hunted and the other not.  This  would entail the creation of a benchmark population (s) somewhere in northern British Columbia as a control.  An intensive, field-oriented study such as  this would do much to reduce some of the uncertainties now facing managers in the Spatsizi.  Moreover, the creation of an effective preserve has other benefits  besides its contribution to harvest policy.  Such a preserve might act as. a  reservoir for the natural restocking of any adjacent, over-exploited game populations.  It could serve as an undisturbed laboratory for other scientific  investigations into wildlife management and, as well, as an attraction to many wilderness users who appreciate being in a wildlife setting where hunting does not take place. The concept, then, can be expressed as a management guideline in the following way:  a wildlife refuge should be established in northern B.C.  for the purpose of serving as a control in the monitoring of hunted wildlife populations elsewhere.  This refuge should encompass, at a minimum, an intact  large mammal, predator-prey system which is neither hunted nor affected'by^any kind of resource development.  Caribou range would likely be one of the criteria  for determining the boundaries of this preserve but much field work coupled with the theoretical criteria for defining wildlife refugia* is required. . An obvious and crucial concern, however, is that the boundaries of a wildlife reserve must correspond to the spatial limits set by the d i s t r i bution and movement of the particular population: i.e. the entire home range. It is pointless to create a protective reserve for any species i f , during * cf. Diamond and May, 1976; Diamond, 1976; Terborgh, 1976; VJhitcomb et a l , 1976: Goeden, 1979: Giles and Scott, 1969; Brown, 1971; Nudds, 1979.  69 seasonal movements, the protected population roams outside the reserve boundaries.  Moreover, i f the protected population is an ungulate species, then  the associated predator populations and their operating territories must be included within the reserve boundaries. noted this requirement  Indeed, Haber (1978) has already  in the case of the Spatsizi.  My central point here is that i f managers cannot effectively guarantee against over-exploitation or other negative impacts on wildlife communities through stringent regulation of hunting activity then the creation of ecologically viable game preserves for the purpose of serving as a benchmark for the study of the effects of hunting on other populations is a potential means of reducing some of the present uncertainties in formulating harvest policies.  70  CHAPTER  VII  Current Wildlife Management Practices in Spatsizi Park - Description and Analysis  7.1  Introduction The purpose of this chapter is to examine current management prac-  tices in Spatsizi Park and assess their consistency with the rules developed in the preceding chapter.  To the extent that current practice correlates with  the desired guidelines, the policy mandate, I contend, is being met. Conversely, deficiencies and inconsistencies between practice and guidelines indicate the degree to which wildlife management in the park fallsjshort of its goals and, therefore, a situation wherein wildlife communities are being :  significantly altered through inadequate or misdirected management. My approach in this'chapter is to.consider each management guideline separately; describing, f i r s t , present practices as they relate to that guideline, assessing the extent to which they are consistent, and finally, identifying the deficiencies in current management approaches.  My conclusions  72 and suggestions about how present practices might be improved to better meet the guidelines are left to the following and final chapter.  7.2  Calculation of the Harvestable Surplus  7.2.1  The Rule To calculate the biological surplus for each population, the  following information is necessary:  population size and age/sex distribution;  net annual increment; densities of important predators and their alternate prey; area! extent of home range; determination of the range carrying capacity, behavioural responses of surviving populations to hunting harassment and loss of herd experiential knowledge. 7.2.2  Present Regulation of the Harvest In December of 1975 when Spatsizi Plateau Wilderness Park was  created, the area's wildlife endowment was subject to the same management policies that prevailed throughout the remainder of British Columbia.  With a  few individual exceptions, provincial wildlife management consisted of two components:  a limited season for hunting and; a limit on the number of animals  taken by each hunter.  With minor variation the hunting seasons for big  game comprised approximately two to three months in the f a l l although a short spring season was allowed in some areas for grizzly bear.  During such  periods the harvest of game animals was permitted by anyone of legal age i  who had purchased a hunting license from the Crown. There were n o i limits i  set as to the number of animals that could be taken from a particular region i  nor were there limits on the number of hunting licenses issued. Managers apparently believed that by restricting hunting activity to two or three months of the year they could successfully regulate the harvest and ensure protection of an adequate breeding stock.  73  Gradually over the years, as over-exploitation trends in some parts of the province were recognized, other devices were implemented to further regulate hunting.  'Quotas' were established from commercial guide/outfitters  and, more recently, 'limited entry hunting' has been introduced for resident hunters in some areas.  Additionally, restrictions on size, age, and sex of  harvestable specimens were emplaced; 'no shooting' areas and 'no hunting  1  areas  were created. In 1975, however, Spatsizi Park was subject to no regulation other than a restricted season and a 'bag limit  1  for hunters.  The establishment of the area as a provincial park brought i t under the jurisdiction of the provincial Parks Branch.  The Order-in-Council stipu-  lated that while the Parks Branch was to be o f f i c i a l l y responsible for wildlife management in the park, biological information and technical advice could be supplied by the Fish and Wildlife Branch.  Since that time harvest limits  have been set by Fish and Wildlife biologists although very recently this has been a joint effort between Parks and Fish and Wildlife personnel. At the time the park was created an ecological reserve at Gladys Lake in the center of the park was established, primarily for Stone's sheep and goat winter range. Branch  This reserve,administered by the Ecological Reserve  cf the Ministry of Environment, was closed to all hunting. ]Addii •;  tionallv, shortened seasons for goat and grizzly bear were imposed on the remainder of the park for the 1975/76 season.  Compulsory reporting, which  does not limit harvests but only increases harvest information for managers, was instituted for sheep and grizzly also in 1975. In 1976 the season for; mule deer and grizzly in Spatsizi Was closed and the seasons for goat, moose, sheep and caribou shortened; pulsory reporting for goat was also commenced.  Com-  74 In 1977, for the f i r s t time, quotas and limited entry hunting for caribou, sheep, goat and grizzly were imposed upon non-resident and resident hunters respectively.  In addition, the recognition that substantial in-  creases in illegal hunting during and subsequent to the construction of the B.C. Railway grade* resulted in restrictions to access along the western perimeter of the park. For the 1980/81 season moose also became the subject of quotas and limited entry hunts.  At present, quotas for black bear and wolf are being  planned. While i t is clear that hunting activity in Spatsizi has been increasingly restricted over the past few years, i t is important for our purpose here to understand what information was used by managers to develop these regulations.  Since 1976, wildlife harvests in Spatsizi have been based pri-  marily, and in some cases solely, on past harvest trends.  Traditional yields  from pre-1976 seasons have been used to determine a percentage harvest rate that is then applied to estimates of total population.  Estimates of total  species populations have, indeed, been made for Spatsizi.  The area, in  fact, has probably had more inventory work carried out than any other in northern British Columbia of similar size.  From 1962 to the present, i t has  had at least 33 aerial surveys carried out over most or portions of the area. Much of the focus was on caribou but since 1977 other resident species have also been surveyed.  Inventory information garnered through aerial surveys  is shown in Table 7.1.  Estimates by park wildlife managers based on:observed f  numbers are also illustrated.  Figures for estimated populations are. calcu-  lated using other information in conjunction with observed counts. /This information includes hunter sightings, hunter sample questionnaires jand harvest data. *  At least 50 caribou during this period according to Bergerud (1978a, p.33).  Table 7.1:  Large Mammal populations in Spatsizi Park. Max. observed on any one survey  moose caribou  216 2573  Tot. population est. by managers 1000-•1300 3000 ± 200*  mule deer  -  20 - 25  mountain goat  163  500 • 700  mountain sheep  294  500 - 600  grizzly bear  4  90 - 100  wolf  78  80 - 100  Figures from unpublished tables prepared by B.C. Parks Br.  * van Drimmelin estimates 1400 in 1981 letter to Flaine Rivers, B.C. Ombudsman's Office.  Uncertainties can arise in direct observation in two ways;' some animals doubtlessly escape observation while others can be double-counted. Methods of estimating total populations derived from direct sightings are subject to error and the biases of individuals doing the calculations.  Here,  for example, a factor of 5 to 6 has been used to obtain the present estimated population of moose although "there have been no intensive moose surveys in the Spatsizi" (Spalding, 1980).  Possibly this figure came from adjustments  made to a 1974/75 suggestion by a guide-outfitter that there were some 2,300 resident moose in the area (Haber, 1978, p.59).  In a memo documenting un-  gulate populations in the Skeena Region (which includes Spatsizi Park) the former regional wildlife biologist for the Fish and Wildlife Branch stated "we have no well documented assessment of (moose) numbers, but are aware, subjectively, of local populations which are increasing, some of which are decreasing, and others which appear stable.  Habitat quality varies among  these areas but habitat assessments, too, are largely subjective. Much more work is needed and annual systematic counts are imperative"(Hatler, 1978). The memo makes an even broader statement regarding a l l ungulates in northwestern British Columbia:  "we have data from too few areas and too few years  to provide solid documentation for population trends." In the past year or so some additional biological information in the form of cow:calf ratios and recruitment has been subjectively considered in the calculations, at least for caribou.  For the 1981/82 hunting seasonj  park wildlife managers have attempted to make caribou harvests more conservative by adopting two measures.  First, the determination of total population i • •-{,  has been based on total observed- animals and not on estimated totals.  Second,  recent harvest rates for caribou set at 10% have been further reduced to 5%. i  By incorporating these measures into their calculations of harvestable surpluses, managers believe that they are well within the limits of sustainable  77 yields.  The problem remains however, as to how to accurately determine a  10% or 5% harvest.  As one of the park wildlife managers said, "to attempt  to regulate sustained yields of 5 or 10 percent of a wildlife population when total inventory information is accurate to only - 50 percent is questionable at best!" (van Drimmelin, pers. comm.).  7.2.3  Deficiencies in Current Practices of Establishing Harvest Levels Current practices of establishing harvest levels in Spatsizi f a l l  short of the management guideline in almost every respect.  Data pertaining  to population size, age/sex distribution, predator densities, and net annual increments are non-existent, incomplete or irregularly obtained and not systematically employed in the setting of harvest limits.  The other information  required by the guideline such as home range size, carrying capacity, and the effects of.hunting harassment on surviving populations is non-existent. Moreover, as I have shown in the preceding section, what population data exists cannot be considered reliable. It is worthwhile at this point to describe the effect past and present management has had on Spatsizi wildlife populations i f only to provide empirical support that the approach indicated by my derived guideline is valid. While the earlier statement by Hatler about the paucity of population data underlines the speculative nature of conclusions drawn about Spatsizi wildlife trends, at least two observers believe that the number of some game populations has declined significantly in the last six years or so.  According to Bergerud  and Butler (1978) Tomias Mountain caribou, the largest of two main herds in the Spatsizi, have declined some 45% in the period 1973-1978.  They speculate that  a recruitment rate of 5-6% has failed to offset a natural mortalityi rate of 12% plus the additional effects of an estimated 3% harvest rate. indicate an annual decline of 9%.  These figures  The second major population, the Caribou  Mountain/Brock Mountain herd, is estimated by them also to have declined approximately 40% during this same period (Figure 7.2).  These  78  1  LUCKHURST  800 700  A  SINCLAIR  o  C0LLINGW00D  +  PRCV. FIRE  600  n  500 -  ®  COUNTS FLATS  HARPER BRUNS  UJ  <  400-  o  3 3  SLOPE OF  300  9%  200 BROCK MT.  100  —I  1972  1  1973  1  1974  /  1  1975  1  1976  1—-  !977  Figure 7.2: Population trends of caribou in Spatsizi Park, (source Bergerud & Butler, 1978)  79 trends are verified in a general way by Haber (1978) in his work on the wolfungulate system in the Spatsizi. Similarly, for moose Bergerud and Butler calculate 9% decline in the 1973-1978 period.  Again, Haber concurs with their conclusion that the Spat-  sizi moose population may be in trouble. Haber (1978) also carried out some preliminary investigation of the status of resident sheep and goat populations.  The former are extremely hard  to inventory and Haber cautions about interpreting the data.  He concludes  that total numbers have remained relatively stable over recent years but:that trophy hunting has severely reduced the proportion of adult males in the population. "There has been a major decrease in the percentage of older males since the onset of heavy resident trophy hunting in 1972, with the resulting possibil i t y of future adverse impacts on other sex and age classes." (Haber, 1978) His available data on goats did not allow him to make any interpretation of population trends in this species. If some big game populations are in decline what does this imply with respect to the way in which harvests are being managed?  Both Bergerud  and Butler (1978) and Haber (1978) have stated that without a temporary suspension of hunting, particularly for moose and caribou, these trends will continue and populations will become seriously and, possibly, irreversibly depleted.  By their calculations there is no harvestable surplus for moose  or caribou at the present time.  :  "The sustained yield of moose and caribou depends on calf survival. At the present time, neither species has any surplus available for harvest - both species are declining. .. .The harvest of caribou is small, but i t adds to the rate of decline. The biological answer to the hunting question is easy - there are no surpluses available for either moose or caribou, and hunting should cease." (Bergerud & Butler, 1978, p.44)  "Pending the results of surveys to determine densities of the moose and caribou populations in particular, a l l moose, caribou, sheep and goat hunting should be stopped at least temporarily in the upper Stikine-Spatsizi region." (Haber, 1978,  p.7)  Two observations can be drawn from the above facts.  First,  park wildlife managers did not employ the necessary biological information to accurately and safely determine sustainable harvests.  Second, when the  results of independent field studies were submitted to them in 1978,  the  recommendations of the researchers were not incorporated into park harvesting policies for the subsequent years.  While the data used by Bergerud arid Haber  were far from perfect, they were nontheless the only comprehensive surveys of Spatsizi done to date. In the f i r s t instance, i t is clear that utilizing mainly traditional hunter harvest trends, with wide-ranging  estimates of total population,  limited recruitment information, and subjectively determined 'safety margins', managers have failed to prevent a depletion in, or alteration to, ungulate populations (again, with the possible exception of goat) - at least up until 1978.  The harvest is s t i l l being maintained on a l l populations which indi-  cates, i f Bergerud and Haber are correct, that over-exploitation is occurring and population declines are being  perpetuated.  In the second instance, instead of responding to the population declines indicated by the field studies (as the management mandate for the park might have obliged them to do in the face of an apparently altering natural progression), wildlife managers reacted by staging a three hour survey flight to inventory caribou.  One small population was observed for which a  recruitment rate of 13.3 percent was determined.  This turned out to.be  sufficient, evidence for the Fish and Wildlife Branch to maintain the 1979 harvest levels, despite the fact that the survey biologist stated that he was "unsatisfied with (his) figures and felt that (he) may have biased them •upwards" (Hatler, 1979).  81  7.3 Age/Sex Distribution of Populations 7.3.1  The Rule All harvests must be structured so that the natural age/sex  distribution of each game population is preserved. 7.3.2  Present Regulation of the Harvest Table 7.2 illustrates the structure of harvest of big game from  Spatsizi Park for the five year period prior to and including 1980. the focus on trophy hunting is dramatically apparent.  Here,  All five big game  species are hunted almost exclusively for trophy (i.e. male) specimens.* 7.3.3  Consistency with the Guideline Current practice directly contravenes the requirement that  natural age/sex distributions be preserved.  Hunting activity in Spatsizi  clearly is selective for mature male specimens.  The natural structure of  these populations, particularly for caribou, sheep, and moose must be undergoing alteration as a result.  Haber (1978), as noted earlier, has a l -  ready observed"this distortion for sheep in the park.  7.4 Management Units 7.4.1  The Rule Where applicable, harvests must be regulated for discrete popu-  lations: policies cannot apply indiscriminately to a l l populations. 7.4.2  Present Management Approach Spatsizi Park is administered as part of Wildlife Management Unit  6-20.  While quotas and limited entry hunts apply to the park they dp not  apply to the remainder of'the area outside the park. *  Similarly, other  Female mountain goat are taken only because hunters cannot distinguish between the sexes in the f i e l d .  Table 7.2:  Harvest of Big Game from Spatsizi Park 1975 - 1980*  Caribou M F 1975 Resident  31  1976 Resident Non-resident  1977 Resident Non-resident  1978 Resident Non-resident  1979 Resident Non-resident  ,1980 Resident Non-resident  Goat  MF  MF  Moose  Grizzly  M F ' M F  9  23  19  25  39  11  34  40  39  11  35  34  39  Non-resident  Sheep  4  2  23  3 5  5  5  5  9  23  6  5 4  23  6  14  8  1  2 3  30  0  16  2  6 2  20  0  0  2  2  15  0  22  4  10 4  25  5  4  2  11  14  5  4 1  20  4  2  2  17  6  8 3  26  7  3  3  2  0  20  6  12  25  1  8  46  2  0  1  12  0  12  2  1  0 I  0 11  anticipated  1981 Resident Non-resident  * Figures quoted in letter to Elaine Rivers, March 19, 1981 from B. van Drimmelin, Regional Wildlife Biologist, Skeena Region. Figures in i t a l i c s for 1975 & 1976 are those cited by Hazelwood (1976).  management units which adjoin the park boundaries visions for restricting the number of animals taken.  lack proFigure 7.3 illustrates  the relationship between park boundaries and the surrounding game management uni ts. One other form of regulatory unit is superimposed over the wildl i f e management units.  Each licensed guide/outfitter in this area has a  certificated territory in which he retains the sole right to guide nonresident hunters. These guiding territories are depicted in Figure 7.4. The main observation to be noted in Figure 7.5 is that while there is some congruence between park boundaries, guiding territories, and wildlife management units, substantial divergence also exists among the three.  This results in a somewhat confusing mosaic of management units  each with its own set of regulations. tories which touch  Three guide/outfitters* have t e r r i -  upon park boundaries (Hazelwood, 1976) although only  one has by far the majority of the park.  In this last case, quotas are  issued to the certificate holder for that area of his territory within Spatsizi only. is unlimited.  His potential harvest in the remainder of his territory This ied to a situation in the 1979/80 hunting season where-  in the guide/outfitter exceeded his quota and was charged under the Wildlife Act by the Fish & Wildlife Branch.  Charges were dropped when i t could not  be demonstrated that the overharvest (of caribou) had taken place within the park boundaries. Similarly, the territories of some guide/outfitters span two or i.  more wildlife management units, each with varying open seasons on different species (or different sexes for the same species).  As a result of this  overlapping of administrative units, each with its own harvesting regime, there clearly is significant potential for confusion and mismanagement. Collingwood, Savage, Love Bros, (now Fleming).  Figure 7.3: Wildlife Management Units surrounding Spatsizi Park with Gladys Lake Ecological Reserve shown in the center of the Park. Scale 1:2,500,000.  F i g u r e 7.4: Spatsizi outfitter territories boundaries.  Park w i t h Guide/ o v e r l a p p i n g park  Figure 7.5: Management Unit boundaries superimp'osed on guide/outfitter territories and Spatsizi Park.  86  While the boundaries of these units correspond for the most part to natural geographic features they do not necessarily conform to natural ecologic units.  Approximately half of their lengths are drawn along river or  creek valleys with the remainder corresponding to heights of land.  In some  cases, particularly in the case of the park, borders are drawn along some arbitrary elevation contour or consist of a straight line between two points. Any real relationship between management boundaries and the natural habitat or home range of the dominant species is virtually non-existent. The most dramatic example of this is the park i t s e l f . The borders of the Spatsizi were determined through an inter-agency consultative process wherein the interests of each agency were accommodated. This resulted in substantial modification of the boundaries as originally proposed by ecologists and biologists.  Boundaries along the northern edge,  for instance, were altered to exclude potential valuable timber stands in the Stikine and Klappan Valley bottoms, areas also considered to be important caribou winter range.  Along the western edge they were set back one mile from  the B.C.R. grade so as not to prohibit any necessary development ancillary to the railway operation. Similarly, identified timber and high mineral potential areas caused park boundaries to be altered in at least three instances: Spruce Hill on the northeast; Cullivan Creek/MacEwan Creek confluence to the northwest; Kluayetz Creek which forms the northern exposure of the Groundhog coal deposit. i  In addition, access corridors through the southern portion of the park have been reserved for potential mineral development to the east of Spatsizi. 7.4.3  Consistency with the Guideline Here again, there is significant departure of management ;practice  from the requirement indicated by the Order-in-Counci1 that harvest regulation be geared to distinct populations. There has been no identification of popu-  -lation groupings and no definition of home range territories within the park.  Clearly, harvest management units are at variance with natural physio-  graphic and biological divisions.  What has been the effect of this upon the  Spatsizi wildlife resources? First, i t should be apparent that in instances where some populations might be protected by restrictive harvest regulations in the park they become vulnerable to more liberal exploitation i f at some point during the year their natural movements take them outside park boundaries.  This has been an issue  in the case of the Spatsizi caribou herds and, apparently, moose populations. The known winter ranges for both these species l i e outside the park, mostly to the north. The principal wintering grounds for Spatsizi caribou seem to be Brock Mountain, Stikine River Valley, and areas further north into the Kehlechoa, Turnagain and Kechika drainages, a l l areas outside the park.  Needless  to say, quotas and limited entry hunt restrictions no longer apply to these animals once they migrate across park boundaries in mid-late October., approximately one month before hunting season is over. The park boundary also transects the summer post-calving range of a large population of Spatsizi caribou in the upper Kluatantan River where the B.C. Railway grade is located.  Potential concerns include increased access  if the railway goes into operation and impacts arising from development of coal deposits in the area (Hazelwood, 1976).  According to Hazelwood, Mt.  Edozadelly and Lawyers Pass area are also critical rutting and wintering ranges for Spatsizi caribou that l i e outside the protection of the park. Additionally, he has identified important sheep range in the upper Duti 5  1  River  drainage as well as an "extremely important mineral lick" for park'ungulate populations, a l l lying outside the park and a l l requiring special protection. Their original exclusion from park territory was a result of other agency concerns (i.e. Forest Service, Department of Mines).  88 Bergerud and B u t l e r (1978) a l s o make the o b s e r v a t i o n t h a t key areas f o r S p a t s i z i  c a r i b o u i n c l u d i n g Brock M o u n t a i n , Lawyers P a s s , and  F i r e F l a t s , are o u t s i d e the park and t h a t t h i s s i t u a t i o n should be a c r i tical  concern f o r management. Bergerud and B u t l e r ( 1 9 7 8 ) , Haber (1978) and Hazelwood (1976)  all  drew a t t e n t i o n . t o the inadequacy o f p r e s e n t park boundaries  to p r o t e c t  a g a i n s t f u t u r e road development i n t o some o f the above i m p o r t a n t c a r i b o u habitat.  Because park b o r d e r s do not conform w i t h the home range t e r r i t o r y  of S p a t s i z i  c a r i b o u , proposed developments such as the Omineca M i n i n g  Road,  the Kutcho Creek Access r o a d , and t h e p o t e n t i a l c o r r i d o r r o u t e through the s o u t h e r n end o f the park t o the B.C.R., g r e a t l y t h r e a t e n c a r i b o u of the i n c r e a s e d hunter access  they w i l l  because  facilitate.  The second major p o i n t t o be made w i t h r e s p e c t t o the p a t t e r n of the h a r v e s t i s t h a t b l a n k e t p o l i c i e s f o r t h e park p r e v a i l .  Population  e s t i m a t e s , r e c r u i t m e n t data (where used a t a l l ) and, most i m p o r t a n t l y , hunter h a r v e s t t r e n d s are a l l aggregate f i g u r e s f o r the park so t h a t i n f o r m a t i o n on a l o c a l watershed o r d i s c r e t e herd b a s i s i s never o b t a i n e d . aggregated  Without d i s -  i n f o r m a t i o n r e g u l a t e d h a r v e s t s , even i f c o n s e r v a t i v e , can l e a d  undetected to o v e r - e x p l o i t a t i o n o f l o c a l  populations.  These two s p a t i a l  c h a r a c t e r i s t i c s of h a r v e s t management, the n o n - c o n f o r m i t y o f  regulatory  u n i t s w i t h e c o l o g i c a l ones and the aggregated h a r v e s t l e v e l s e s t a b l i s h e d , l e a d a g a i n to the c o n c l u s i o n t h a t management o f park w i l d l i f e w i l l  not  achieve p o l i c y o b j e c t i v e s .  7.5 7.5.1  P u l s e -Harvesting The Rule H a r v e s t s must not be s e t a t f i x e d l e v e l s but must vary from season  t o season as c o n d i t i o n s p e r m i t .  89 7.5.2  Present Practice Allowable harvest levels in Spatsizi, as noted in Section 7.2.2., have  been progressively reduced from unlimited (pre-1976) up to levels "estimated" to be 5% of the total population (e.g.: caribou for '81 season). Note, however, the objective apparently has always been to arrive at some satisfactory or 'optimum' level of harvest that is expressed as a fixed percentage of total population.  This tendency is reflected at the pro-  vincial level where harvest objectives are so established (Table 7.3). 7.5.3  Consistency with the Guideline While the gradual reduction of harvest levels for some park species  must be viewed as an increasingly cautious approach being undertaken by park wildlife managers, i t is apparent that they believe that, at some low level, a fixed percentage harvest of total population for each species can be maintained.  Thus, no provision for the dynamic variability of wildlife com-  munities and, in particular, random environmental sing true biological surpluses.  events, is made in asses-  This administrative propensity to stabilize  the yield of wildlife resources at some fixed level over successive generations decreases the resilience of the system by prohibiting population rebound during seasons of adverse conditions; conversely, i t forfeits a greater return to consumptive users when conditions are favourable for high biological production. There is an additional comment worth making here.  If yields from  a resource are fixed at some rate there develops a tendency on the part of both managers and beneficiaries (here, hunters and guide/outfitters) to expect that rate of return into the forseeable future, particularly so i f development capitalization is tied into anticipated economic returns.  This expectation clearly  reduces the f l e x i b i l i t y available to managers to adjust the rate of resource harvest when conditions appear to warrant i t .  The resource harvester perceives  a loss in benefits to himself with the result that managers are commonly placed  Table 7.3. Provincial harvest objectives for big game species. species  deer moose elk cari bou sheep goat grizzly bear black bear  % harvest  rate  12.6 10 13.6 5 3 3 5 5-10  Proposed Wildlife Management Plan for British Columbia. Munro, 1979.  91  under some c o n s i d e r a b l e p r e s s u r e  t o m a i n t a i n the l e v e l o f e x p l o i t a t i o n .  Such e x p e c t a t i o n s o f r e s o u r c e u s e r s would l i k e l y not a r i s e under a system of f l u c t u a t i n g y i e l d s and, hence, p o l i t i c a l  and economic p r e s s u r e s  on the  r e s o u r c e would be moderated.  7.6  Predator  7.6.1  Management  The Rule P r e d a t o r c o n t r o l should not be implemented as a means to r e s t o r e  ungulate 7.6.2  populations.  Present P r a c t i c e W h i l e p r e d a t o r s , p a r t i c u l a r l y w o l v e s , have been viewed by some  managers t o c o n s t i t u t e a problem i n S p a t s i z i  Park w i t h r e s p e c t to p e r c e i v e d  d e c l i n i n g u n g u l a t e p o p u l a t i o n s , c o n t r o l programs to date.  Nonetheless,  t h e r e has been s i g n i f i c a n t p r e s s u r e  hunters and some advocacy from b i o l o g i s t s tore ungulate populations tained.  have not been implemented i n the p a s t from  t h a t w o l f c o n t r o l i s needed to  res-  so t h a t t r a d i t i o n a l h a r v e s t l e v e l s can be m a i n -  Wolf c o n t r o l a p p a r e n t l y has been p r a c t i c e d u n o f f i c i a l l y by g u i d e /  outfitters  i n the area where and when the o p p o r t u n i t y a r o s e .  t o what e x t e n t t h i s p r a c t i c e i s m a i n t a i n e d a t p r e s e n t .  I t i s not known  The F i s h & W i l d l i f e  Branch has a r t i c u l a t e d a p o l i c y of p r e d a t o r c o n t r o l * i n i n s t a n c e s i s proven t h a t wolves are s e r i o u s l y  depressing  other w i l d l i f e  below l e v e l s needed to meet management o b j e c t i v e s . " what u n d e r l a y a 1978 agency i n the S p a t s i z i  This  "where  it  populations  policy is  undoubtedly  i n i t i a t i v e to implement i n t e n s i v e w o l f c o n t r o l  and o t h e r n o r t h e r n r e g i o n s .  The s t i m u l u s  for t h i s ,  apart  from p e r c e i v e d u n g u l a t e d e c l i n e s by g u i d e / o u t f i t t e r s , was i n p a r t a r e s u l t o f • B e r g e r u d and B u t l e r ' s *  r e s e a r c h i n the S p a t s i z i .  P r e l i m i n a r y Wolf Management  They concluded t h a t low  P l a n f o r B r i t i s h C o l u m b i a , Oct. 1979,  p.ii.  caribou recruitment was caused by high predation of calves and that populations were declining naturally as a result (Bergerud and Butler, 1978).  Hunting,  according to them, would never be possible on a sustained yield basis unless predators were sufficiently controlled. It was this intention, at least in part, that prompted managers to contemplate wolf control for areas in northern British Columbia, including the Spatsizi where apparent ungulate declines were being observed. Bergerud and Butler's conclusion must be examined in light of the multiple equilibrium model which indicates that there are population-densities at which predation is not a limiting factor and a harvestable surplus exists (Haber, 1978).  Predation apparently only becomes a problem when hunting has  pushed ungulate populations close to the escape threshold. cautions against the dangers of this situation:  "  Haber (1978)  only a year or two  of further human harvest, even at a low rate, could accelerate the decline and push the population below the threshold where increments become negative" - Haber, 1978, p.64 (emphasis mine).  Below the escape threshold  natural predation alone could then exert a depensatory pressure on Spatsizi ungulates, even i f harvests are suspended (Haber, 1976, 1977).  Low popu-  lations of some species would then exert higher predation pressures on any alternate prey that were abundant resulting in smaller annual increments and higher threshold densities at which populations enter an irreversible decline (Haber, 1978). Haber also argues that low density caribou populations may be the norm in Spatsizi.  Predator switching by wolves feeding on healthy moose popu-  lations could have suppressed low caribou numbers since the initial colonization.  He finds no evidence in any of the information generated from Spatsizi  research which supports the need for predator control.  Despite these argu-  «•  ments, the general situation perceived by local people, hunters, and some  agency personnel  (Parks Branch  and F i s h & W i l d l i f e ) i s t h a t wolf  are r e l a t i v e l y high while ungulate  populations a r e low.  d i t i o n s , t h e r e f o r e , a r e such t h a t here  populations  The p o l i t i c a l  con-  i s a continual, i f not well publicized,  lobby on t h e p r o v i n c i a l government t o i n i t i a t e wolf c o n t r o l i n t h e north. 7.6.3  Consistency with the Guideline The  f a c t that predators are not being c o n t r o l l e d i n the area a t  p r e s e n t means t h a t t h i s g u i d e l i n e i s n o t b e i n g v i o l a t e d . p r e d a t o r c o n t r o l e v e r be implemented lations a v a i l a b l e to hunters vention of the  as a means o f i n c r e a s i n g u n g u l a t e  i n t h e park, then t h i s must be viewed  predator c o n t r o l might  be c o n s i d e r e d a p p r o p r i a t e .  adequate  f i e l d documentation  human a c t i v i t y u p o n S p a t s i z i u n g u l a t e the years to reduce  arise  T h i s s i t u a t i o n would  t h a t the impacts  p o p u l a t i o n s had been s u f f i c i e n t  of over  p o p u l a t i o n s t o l e v e l s where p r e d a t i o n becomes a c o n t r o l l i n g  ( i . e . a s h i f t from high s t a b i l i t y domain t o l o w ) . In t h i s case, a  d e c i s i o n might and  popu-  as a c o n t r a -  e a r l i e r , i t i s p o s s i b l e that a s i t u a t i o n might  be t h e d i s c o v e r y , t h r o u g h  factor  however,  Order-in-Counci1.  As m e n t i o n e d wherein  Should,  here  be made t o r e s t o r e u n g u l a t e  i t might  ungulates  be n e c e s s a r y  can increase  populations to historical levels  to temporarily reduce  predator levels until  sufficiently.  However, and as I have c a u t i o n e d  i n Chapter  m u s t be c a r e f u l l y made a n d i n l i g h t o f s e v e r a l i m p o r t a n t F i r s t , a better understanding  V I , such a d e c i s i o n bits of information.  of the causal mechanisms o f ungulate  declines  must be a t t a i n e d as w e l l as t h e n a t u r a l d y n a m i c s o f t h e p r e d a t o r - u n g u l a t e system  in question.  is c r u c i a l  P o p u l a t i o n trend i n f o r m a t i o n f o r both predators and prey  so t h a t t h e d e c l i n e as w e l l as h i s t o r i c a l  d e n s i t i e s c a n be  mented and, t h e r e f o r e , the s h i f t i n s t a b i l i t y domains s u b s t a n t i a t e d .  docuFurther-  m o r e , t h e p o p u l a t i o n d e n s i t i e s o f a l t e r n a t e p r e y m u s t be known s o t h a t v i a b l e  94  s t r a t e g i e s o f p r e d a t o r c o n t r o l can be d e v e l o p e d .  F i n a l l y , the obvious  policy  r e q u i r e m e n t i n a s i t u a t i o n of t h i s k i n d i s t h a t h u n t i n g must a l s o be t e r m i n a t e d d u r i n g the p e r i o d of r e c o v e r y .  7.7  The E s t a b l i s h m e n t o f Benchmark  7.7.1  Populations  The Rule In o r d e r t o reduce u n c e r t a i n t i e s a r i s i n g from the e s t a b l i s h m e n t o f  h a r v e s t l e v e l s , benchmark p o p u l a t i o n s must be c r e a t e d f o r the m o n i t o r i n g o f t h e e f f e c t s o f h u n t i n g on l a r g e mammal p r e d a t o r - p r e y systems found i n S p a t s i z i .  7.7.2  Present P r a c t i c e As noted p r e v i o u s l y , when S p a t s i z i  Plateau Wilderness  Park was  c r e a t e d , an e c o l o g i c a l r e s e r v e a t Gladys Lake i n the m i d d l e o f the park was also established.  This  ' e c o r e s e r v e ' was s e t a s i d e t o p r e s e r v e s e v e r a l  unique  p l a n t communities o f the i n t e r i o r s u b a l p i n e zone and a l p i n e t u n d r a as w e l l a r e s i d e n t p o p u l a t i o n o f S t o n e ' s sheep. serves  A d d i t i o n a l l y , the r e s e r v e  as a p r o t e c t e d , i m p o r t a n t w i n t e r i n g range f o r n e i g h b o u r i n g  o f sheep, g o a t s , moose, and c a r i b o u . Lake E c o l o g i c a l Reserve.  as  also populations  No h u n t i n g i s p e r m i t t e d i n the Gladys  A s i d e from t h i s e c o r e s e r v e t h e r e are no o t h e r  t a b l i s h e d w i l d l i f e r e f u g i a i n the S p a t s i z i and upper S t i k i n e a r e a .  es-  All  animal p o p u l a t i o n s w i t h i n the upper S t i k i n e watershed and a d j a c e n t r i v e r basins" i n n o r t h e r n B r i t i s h Columbia are s u b j e c t to h u n t i n g 7.7.3  activity.  C o n s i s t e n c y w i t h the G u i d e l i n e I t might be argued t h a t having the one w i l d l i f e r e s e r v e a t Gladys  Lake i s a l l t h a t i s necessary  to p r o t e c t an adequate p o p u l a t i o n o f  w i l d l i f e s p e c i e s and, therefore,;, s a t i s f y the d e s i r e d g u i d e l i n e .  resident  This  ques-  t i o n can be r e s o l v e d by examining the e x t e n t t o which the Gladys Lake r e s e r v e  protects the various big game populations.  The vital criterion for such a  refuge is that its boundaries conform, at a minimum, to the outer limits of the particular population's home range.  There must be no regular trans-  gression of borders by either the protected animals (unless by natural dispersion) or by hunters.  With respect to Stone's sheep, the degree of  protection offered by the boundaries of the ecoreserve has been called into question.  Hazelwood (1976) has noted that the ecoreserve borders are drawn  along heights of land and, consequently, have l i t t l e meaning for high elevation species such as sheep and goat whose regular movements cover both sides of a mountain.  According to Hazelwood, drainage courses define more  appropriate and natural breaks in the habitat of these species.  It i s ,  therefore, not too improbable to assume that Gladys Lake sheep are exposed a significant amount of the time to hunters operating along the perimeters of the reserve boundaries. Second, the matter of regulating and enforcing a prohibition on hunting in the reserve is a major concern.  As recently as the 1979/80 season  there were no signs posted at Coldfish Lake (the principal entry point for the Gladys Lake Ecological Reserve) to warn hunters of the protected area and hunters there were making inquiries as to what the hunting opportunities were like in the area covered by the reserve.  There is no regular monitoring  of hunter activities in the Gladys Lake area or the park generally. The important observation that can be made.about this ecoreserve, however, is that i t does not protect those populations, principally moose and caribou, that seem to be in the most trouble. There is no preserve in the upper Stikine (or anywhere else in British Columbia) set aside for this purpose. Every population of caribou and moose in the Spatsizi territory appears to be vulnerable to hunting at some point during the annual hunting season. final point has to be made about the establishment of such a preserve.  One  96 W h i l e i t c o u l d be argued t h a t i t i s s u f f i c e n t t o , d e l i n e a t e p r e s e r v e  boun-  d a r i e s o n l y so t h a t the p o p u l a t i o n i s p r o t e c t e d d u r i n g h u n t i n g s e a s o n , a p r e s e r v e so d e f i n e d would not be e f f e c t i v e i n p r o t e c t i n g the p o p u l a t i o n and, perhaps more i m p o r t a n t l y , i t s h a b i t a t from o t h e r k i n d s o f human a c t i v i t y and encroachment t h a t are bound to ensue a t some f u t u r e p o i n t i n the upper S t i k i n e . Thus, the p r e s e r v e must embrace the e n t i r e range o c c u p i e d by the p o p u l a t i o n d u r i n g the c o u r s e o f an annual c y c l e . T h i r d , no attempt i s being made t o m o n i t o r the impacts o f on any p r o v i n c i a l w i l d l i f e p o p u l a t i o n by the government. Ecological  Even i f Gladys  Reserve was an e f f e c t i v e w i l d l i f e r e s e r v e , i t s u s e f u l n e s s  p o s s i b l e c o n t r o l t o measure h u n t i n g impacts i s not being  hunting  utilized.  as a  Lake  C H A P T E R  VIII  C o n c l u s i o n s and Recommendations  8.1  Synthesi s In the f o r g o i n g I have attempted  t o i d e n t i f y the s a l i e n t f e a t u r e s  o f p r e v a i l i n g w i l d l i f e management i n S p a t s i z i P l a t e a u W i l d e r n e s s Park and to a s s e s s t h e s e w i t h r e s p e c t to t h e i r c o n s i s t e n c y w i t h the O r d e r - i n - C o u n c i 1 . have done t h i s by i n t e r p r e t i n g the p o l i c y s t a t e m e n t  i n the  I  Order-in-Counci1  i n terms o f i t s i m p l i c a t i o n s f o r the management o f n a t u r a l systems and,  from  t h i s i n t e r p r e t a t i o n , have d e v e l o p e d a s e t o f management g u i d e l i n e s t h a t s h o u l d be a p p l i e d t o the a d m i n i s t r a t i o n o f park w i l d l i f e r e s o u r c e s i f g o a l s are t o be met.  These g u i d e l i n e s , d e r i v e d from f u n d a m e n t a l e c o l o g i c a l p r i n -  c i p l e s , were then used as c r i t e r i a by w h i c h t o a s s e s s c u r r e n t management practi ces. Of the s i x r u l e s so d e v e l o p e d  o n l y one, the one r e l a t e d to the pro-  h i b i t i o n of p r e d a t o r c o n t r o l , can be s a i d t o be i n c o r p o r a t e d i n t o p r e s e n t  98  park policy.  The remaining five are contravened by current practices  (Table 8.1).  In summary, the information employed to set present harvest  levels primarily includes past harvest information, in some instances subjectively coupled with widely ranging estimates of total population. Regular, accurate surveys of species populations, age/sex distribution, and recruitment are not conducted.  No information pertaining  to home range size, carrying capacity, or species behavioural responses to hunting harassment has been developed or used in establishing harvests. While over the past two to three years park wildlife managers have set permissible harvest levels and imposed restrictions on park hunting activity, the levels so established tend to be based on some pre-determined  percentage  yield objective rather than a fluctuating level based on prevailing conditions. Harvest levels also are established for park populations as a whole without any reference to what discrete populations might exist and what home ranges might transgress park boundaries.  And most importantly, perhaps, the  hunting activity within the park is almost exclusively trophy hunting which implies that the natural age/sex distribution of the hunted populations is being altered, not to mention possible other more intangible and  long-term  impacts. Finally, no viable wildlife reserves have been established to ensure maintenance of breeding populations of big game species found in the upper Stikine.  No other means of minimizing uncertainties in the de-  termination of harvest levels have been substituted for the reserve concept. I conclude here that current wildlife management practices in Spatsizi Park result in substantial risks that the mandate laid out in the Order-in-Councii may well be violated.  That is to say, current regulation  of big game harvests is such that the preservation of natural communities and the unaltered progression of ecological systems is quite unlikely to be achieved.  T a b l e 6."!. C o n f o r m i t y  o f S p a t s i z i management p r a c t i c e s w i t h d e r i v e d management g u i d e l i n e s Derived  Management  Practiced  Guidelines  Non-linearities equi1ibria  Dynamic  3. 4.  & mulitiple  variability  Inter-connectedness Uncertainties  Spatsizi  • 1 .  Harvestable surpluses based on population density, recruitment, age/sex d i s t r i b u t i o n , predator and a l t e r n a t e p r e y d e n s i t i e s ;  No  b.  Pulse  No  harvesting;  Harvests managed f o r d i s c r e t e populations;  No  4.  Predators  Yes  5.  Age/sex d i s t r i b u t i o n  -6.  in  not controlled; maintained;  Wildlife refuge established  No No  U3  100  In f a c t , the apparent d e c l i n e of several ungulate populations in the park lends a special urgency to t h i s conclusion.  If p o l i c y makers are serious about  the goal of maintaining the i n t e g r i t y of natural communities then i t may be that a r e - e v a l u a t i o n of the hunting provision in the Order-in-Counci1 i s necessary.  My point here i s that i t i s probable there i s a fundamental  c o n t r a d i c t i o n in the p o l i c y statement in that the stated goals are incompatible.  I believe t h i s i s impossible to prove with the a v a i l a b l e evidence;  much more research i s required to resolve the question.  Under the assump-  t i o n that the p r e s e r v a t i o n i s t i n t e n t of the stated p o l i c y i s the overr i d i n g concept, however, I argue here that the onus i s on park w i l d l i f e managers to s a t i s f a c t o r i l y demonstrate the c o m p a t i b i l i t y of hunting, sustained y i e l d or otherwise, with the preservation of i n t e g r i t y of the w i l d e r ness ecosystem before any hunting be permitted.  8.2  Recommendations Park w i l d l i f e managers have two courses of a c t i o n open to them i f  consistency i s to be achieved between management o b j e c t i v e s and present p r a c t i c e in S p a t s i z i Park.  The f i r s t i s to s u b s t a n t i a l l y increase f i e l d  research so that the necessary b i o l o g i c a l information can be gathered and incorporated into harvest p o l i c i e s .  This w i l l require a serious  funding  commitment by the government that i s u n l i k e l y to happen in the near f u t u r e . Even i f funding requirements were s a t i s f i e d there would be s u b s t a n t i a l d i f f i c u l t y in adhering to the guidelines because (a) of the problem of protecting animals moving in and out of the park; and (b) of the problem of reaching agreement amongst the various agencies to e s t a b l i s h the proposed reserve.  101  The second a l t e r n a t i v e open to government i s to d e s i g n a t e S p a t s i z i Park t h e c o r e o f t h e p r o p o s e d r e s e r v e and expand i t s b o u n d a r i e s so t h e y conform t o t h e a p p r o p r i a t e t e r r i t o r i e s o f park w i l d l i f e p o p u l a t i o n s . r e s e r v e c o u l d s e r v e a s a benchmark f o r a l l o f n o r t h e r n B r i t i s h  Such a  Columbia.  The government has n o t p u r s u e d , and does n o t a p p e a r e a g e r t o a d o p t , e i t h e r o f t h e s e two a l t e r n a t i v e s . While t h i s s t u d y may n o t e f f e c t changes i n government p o l i c y f o r S p a t s i z i , I f e e l i t a p p r o p r i a t e t o recommend a s p e c i f i c c o u r s e o f a c t i o n t h a t embodies elements o f both t h e o p t i o n s o u t l i n e d above. This course includes: 1.  A temporary  s u s p e n s i o n o f a l l h u n t i n g a c t i v i t y w i t h i n t h e park  and w i t h i n t h o s e a r e a s a d j a c e n t t o t h e park p r e s e n t l y b e l i e v e d to c o r r e s p o n d t o t h e home range t e r r i t o r y o f any S p a t s i z i w i l d l i f e populations, particularly caribou. 2.  The e x e c u t i o n o f s t u d i e s t o d e v e l o p t h e n e c e s s a r y i n f o r m a t i o n for:  a) t h e d e l i n e a t i o n o f w i l d l i f e p o p u l a t i o n s and home range  t e r r i t o r y f o r each p o p u l a t i o n u t i l i z i n g S p a t s i z i Park f o r p a r t o r a l l o f i t s s e a s o n a l movements;  b) annual s u r v e y s on an on-  g o i n g b a s i s o f p o p u l a t i o n s numbers, age/sex d i s t r i b u t i o n , r e c r u i t m e n t , range c o n d i t i o n ; c) a d e t a i l e d u n d e r s t a n d i n g o f t h e dynamics o f r e s i d e n t p r e d a t o r - p r e y systems; and d) a more ccm-prehensive  u n d e r s t a n d i n g o f t h e impacts o f h u n t i n g upon  the b e h a v i o u r a l r e s p o n s e s o f hunted p o p u l a t i o n s . 3.  An a d j u s t m e n t o f t h e b o u n d a r i e s o f the G l a d y s Lake E c o l o g i c a l R e s e r v e so t h a t  they  conform t o t h e known range o f t h e Gladys  Lake sheep p o p u l a t i o n . * * As t h i s f i n a l d r a u g h t i s b e i n g p r e p a r e d t h e w r i t e r has beer: i n f o r m e d by G. Hazclv.'ocd (pers.comm.) t h a t t h e b o u n d a r i e s o f t h i s e c o l o g i c a l r e s e r v e have, i n f a c t , been changed j u s t r e c e n t l y t o c o r r e c t f o r t h i s . I t i s p r o b a b l y t o o e a r l y t o a s s e s s t h e e f f e c t i v e n e s s o f t h i s change b u t from the d e s c r i p t i o n I r e c e i v e d i t sounds as i f the major d e f i c i e n c i e s have been a m e l i o r a t e d .  102 4.  The conduct of a study to d e v e l o p boundary c r i t e r i a f o r the s e l e c t i o n of a w i l d l i f e r e f u g e i n n o r t h e r n B r i t i s h Columbia w h i c h encompasses a t a minimum, the complete home range  terri-  t o r y o f a l a r g e mammal p r e d a t o r - p r e y system w i t h a l l i t s c o n s t i t u e n t p o p u l a t i o n s of the type found i n S p a t s i z i 5.  Park.  The s t a t u t o r y c r e a t i o n o f a w i l d l i f e r e f u g e conforming t o the a r e a d e f i n e d above and having the same k i n d o f use r e s t r i c t i o n s as an e c o r e s e r v e  ( i . e . no h u n t i n g , no development, minimal  i n t e r f e r e n c e by man). Some comments on these recommendations a r e p e r t i n e n t . i s s e l f - e x p l a n a t o r y and f o l l o w s l o g i c a l l y from the p r e c e d i n g  The  first  discussion:  s e v e r a l r e s i d e n t s p e c i e s are a p p a r e n t l y being reduced i n a b s o l u t e numbers and/ o r a r e beimg a l t e r e d w i t h r e s p e c t to age/sex s t r u c t u r e o f t h e i r p o p u l a t i o n s . Managers do not have the i n f o r m a t i o n n e c e s s a r y to i n s t i t u t e sound a m e l i o r a t i v e p o l i c i e s t o c o u n t e r a c t t h e s e t r e n d s and, t h e r e f o r e , c o n t i n u e d h u n t i n g o n l y e x a c e r b a t e t h e s e problems and t h w a r t the achievement o f p o l i c y The second recommendation focuses on the need f o r park  can  goals.  wildlife  managers to c a r r y out s u b s t a n t i v e f i e l d r e s e a r c h b e f o r e an a p p r o p r i a t e l e v e l o f h u n t i n g a c t i v i t y , i f such e x i s t s , can be soundly e s t a b l i s h e d f o r w i l d l i f e populations.  This raises  another p o i n t .  Spatsizi  Managers commonly c l a i m  t h a t the reason they cannot g a t h e r s u f f i c i e n t i n f o r m a t i o n t o e f f e c t i v e l y manage the r e s o u r c e i s a l a c k o f f u n d s .  The i m p o r t a n t q u e s t i o n f o r managers then  becomes one o f what to do g i v e n inadequate monies to do a p r o p e r j o b . p r o p e n s i t y i n B r i t i s h Columbia has been to carry  on a d m i n i s t e r i n g the  The harvest  w i t h minimal o r no d a t a , and on the assumption t h a t p a s t h a r v e s t l e v e l s were sustainable.  T r a d i t i o n , i t seems, has played an u n s u p p o r t a b l y prominent r o l e  i n the s e t t i n g out o f management p r i o r i t i e s .  However, the absence o f adequate  f u n d i n g i s no excuse t o p e r p e t u a t e a p r a c t i c e t h a t appears t o be r u n n i n g a  103 serious  r i s k of contravening p u b l i c p o l i c y o b j e c t i v e s . The l a s t t h r e e recommendations a l l f o c u s on a c r u c i a l  issue.  Nowhere i n t h i s p r o v i n c e has t h e r e been e x p r e s s l y s e t a s i d e a t r a c t o f w i l d l a n d f o r the p r o t e c t i o n and p r e s e r v a t i o n o f r e p r e s e n t a t i v e b i g game popul a t i o n s c o - e x i s t i n g as a n a t u r a l , u n a l t e r e d b i o l o g i c a l community. few a r e a s o f r e p r e s e n t a t i v e h a b i t a t have been p r e s e r v e d national  ( p r i n c i p a l l y the  p a r k s ) most b i g game p o p u l a t i o n s i n B r i t i s h Columbia a r e exposed to  h u n t i n g through n a t u r a l seasonal protected  movements which b r i n g them o u t s i d e o f  zones. Two p o i n t s a r i s e as t o the i m p l e m e n t a t i o n o f w i l d l i f e  First,  While a  i t i s c o n c e i v a b l e t h a t any p e r m i t t e d l e v e l o f h u n t i n g i s  w i t h the p r e s e r v a t i o n o f n a t u r a l systems.  reserves. incompatible  In such a case the o n l y  consistent  management o p t i o n i s t o suspend h u n t i n g o f S p a t s i z i w i l d l i f e and l e t n a t u r a l processes  d e t e r m i n e the c o u r s e o f events i n the a r e a .  T h i s would not a l t e r  the i n t e n t i o n s o f the O r d e r - i n - C o u n c i 1 but o n l y e l i m i n a t e one o f the s e v e r a l " p e r m i t t e d " uses o f the p a r k . Second, w h i l e i t may be t h a t some l e v e l of h a r v e s t i s without, s i g n i f i c a n t l y a l t e r i n g the n a t u r a l systems, biological  possible  such a g r e a t volume o f  d a t a would be r e q u i r e d f o r proper management t h a t one might  r e a s o n a b l y wonder whether the law o f d i m i n i s h i n g r e t u r n s would e l i m i n a t e any b e n e f i t s from the a c t i v i t y .  A t r a d e - o f f e x i s t s here.  i n o r d i n a t e amount o f manpower and f i n a n c i a l r e s o u r c e s  Do we expend an  in a herculean e f f o r t  to f u r t h e r reduce (but never q u i t e e l i m i n a t e ) u n c e r t a i n t y o r do we c r e a t e a r e s e r v e which bans h u n t i n g o f an e c o l o g i c a l l y i n t a c t b i g game community, e l i m i n a t e s the p r i n c i p a l r i s k problem a l t o g e t h e r , and can be executed w i t h a minimal e x p e n d i t u r e o f e f f o r t ? I t would not be a p p r o p r i a t e to end t h i s t h e s i s w i t h o u t some c l o s i n g remarks about my own p e r c e p t i o n s towards S p a t s i z i  Park.  C e r t a i n l y my personal  v a l u e s about the importance o f w i l d ! animal s p e c i e s t o the o v e r a l l w e l l - b e i n g  104 o f n a t u r a l e c o s y s t e m s a n d t o h u m a n e n j o y m e n t l e d me t o s e l e c t w i l d l i f e p o l i c y as a subject f o r i n v e s t i g a t i o n i n the f i r s t place. controversy over hunting  p r a c t i c e s i n S p a t s i z i Park,  The p u b l i c  a s w e l l a s my  v i s i t s t o i t , l e d me t o b e l i e v e t h a t t h i s a r e a w a s a m o r e t h a n choice to pursue  t h e more focused examination  management I wished  to carry out.  the Order-in-Council without ecosystem  I agree  that comprises  of provincial wildlife  f u l l y with the objectives of  natural communities".  S p a t s i z i Park and i t s s u r r o u n d i n g s  far too valuable (as a w i l d l i f e heritage, wilderness of i n s p i r a t i o n a l and educational  enrichment)  The  i s ,I  contend,  laboratory, or  source generations  away by i n e f f e c t i v e  i s overdue f o r a sound r e - o r d e r i n g o f w i l d l i f e  priorities in Spatsizi.  proceed  wilderness  f o r present and f u t u r e  o f B r i t i s h C o l u m b i a s t o be j e o p a r d i z e d o r s q u a n d e r e d The time  adequate  to "allow the progression o f natural systems to  a l t e r a t i o n " and t o "maintain  management.  own  management  105 Bibliography  A l l e n , E.D., J . V . B a s i l e , R.G. J a n s o n , T.N. L o n n e r , L . J . L y o n , C . L . M a v c a n , F. P o n d , R. B e a m , a n d D. S a i l . 1976. 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