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Habitat selection and use in winter by moose in sub-boreal forests of north-central British Columbia,… Eastman, Donald Sidney 1977

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HABITAT SELECTION AND USE IN WINTER BY MOOSE IN SUB-BOREAL FORESTS OF NORTH-CENTRAL BRITISH COLUMBIA, AND RELATIONSHIPS TO FORESTRY by DONALD SIDNEY EASTMAN B.Sc,  U n i v e r s i t y o f B r i t i s h Columbia 1962  M.Sc,  U n i v e r s i t y o f Aberdeen 1964  A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY  in THE FACULTY OF GRADUATE STUDIES Department o f P l a n t  Science  We a c c e p t t h i s t h e s i s -as conforming t o the r e q u i r e d s t a n d a r d  THE UNIVERSITY OF BRITISH COLUMBIA December 1977 (c)  Donald Sidney Eastman  In presenting  t h i s t h e s i s i n p a r t i a l f u l f i l l m e n t of  the r e q u i r e m e n t s f o r an advanced degree a t the U n i v e r s i t y o f B r i t i s h Columbia, I agree t h a t the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and study. that permission f o r extensive  I f u r t h e r agree  copying of t h i s t h e s i s f o r  s c h o l a r l y purposes may be g r a n t e d by the Head o f my Department o r by h i s r e p r e s e n t a t i v e s .  I t i s understood  copying or p u b l i c a t i o n of t h i s t h e s i s f o r f i n a n c i a l s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n .  Department o f P l a n t S c i e n c e The U n i v e r s i t y of B r i t i s h Columbia 2075 Wesbrook P l a c e Vancouver, B.C., Canada V6T 1W5 Date  gain  that  Chairman:  Dr. V. C. B r i n k ABSTRACT  A study o f w i n t e r h a b i t a t s e l e c t i o n and use by moose 2 was conducted  i n a 11,300 km  area of n o r t h - c e n t r a l B r i t i s h  Columbia from May 1971 t o August 19 73.  The study area was  l o c a t e d w i t h i n the f o r e s t e d s u b - b o r e a l spruce b i o g e o c l i m a t i c zone, a zone t h a t i s r e c e i v i n g i n c r e a s e d development, e s p e c i a l l y by f o r e s t r y .  H a b i t a t s e l e c t i o n and use was  examined m a i n l y be p e l l e t group surveys and a e r i a l transects.  W i n t e r i n g moose used p a r t i a l c u t o v e r s and burns  more than c o n i f e r o u s f o r e s t s ; c l e a r c u t s were used l e a s t . p a t t e r n i n summer.  deciduous  f o r e s t s and r e c e n t  L i m i t e d d a t a suggested  a similar  W i n t e r use t y p i c a l l y i n c r e a s e d from n e a r  zero a f t e r a r e c e n t d i s t u r b a n c e such as c l e a r c u t t i n g , t o a peak sometime between 10 and 25 y e a r s l a t e r , then d e c l i n e d to low l e v e l s d u r i n g 25 and 9 0 y e a r s , and then a p p a r e n t l y s t a b i l i z e d i n t h e mature f o r e s t stage a t s l i g h t l y levels.  higher  On one i n t e n s i v e l y surveyed a r e a , moose s e l e c t e d  p a r t i a l c u t o v e r s and creek bottoms even though these h a b i t a t s comprised  l e s s than 6 p e r c e n t o f the a r e a .  Moose  began c o n c e n t r a t i n g on w i n t e r ranges a t l e a s t by midNovember, reached a peak i n November-January, and d e c l i n e d steadily thereafter.  Food h a b i t s and d i e t were examined by  I l l  rumen a n a l y s i s , t r a i l i n g and p o s t - w i n t e r  browse  surveys.  Moose had c a t h o l i c d i e t s b u t a t e p r i m a r i l y deciduous browse f o r most of t h e y e a r . late winter.  S u b a l p i n e f i r becomes i m p o r t a n t i n  Diet v a r i e d according  t o season and h a b i t a t .  P r e f e r r e d s p e c i e s t y p i c a l l y were l e a s t common.  Tagged t w i g  t r a n s e c t s r e v e a l e d t h a t moose f r e q u e n t l y browsed p l a n t s more than once b u t r a r e l y re-browsed a t w i g .  The time o f  b r o w s i n g v a r i e d by s p e c i e s and by h a b i t a t w i t h most use r e c o r d e d i n J a n u a r y and i n A p r i l .  Levels of u t i l i z a t i o n  were a l l l e s s than 100 p e r c e n t o f the p r e v i o u s production.  Utilization  (weight-basis)  year s 1  ranged from 33 p e r -  c e n t on r e d - o s i e r dogwood t o 3 p e r c e n t on s u b a l p i n e  f i r ; and  from t r a c e amounts i n an u p l a n d burn h a b i t a t t o more than 40 p e r c e n t i n deciduous f o r e s t , p a r t i a l c u t o v e r bottom h a b i t a t s .  and r i v e r  Bedding .habits were examined i n an attempt  t o d e f i n e c o v e r f o r moose.  Moose choose upper s l o p e s  that  f a c e d s o u t h p a r t i c u l a r l y when snow depths became r e s t r i c t i v e (> 8 0 cm).  Moose tended t o s e l e c t l a r g e r than average t r e e s  and t o bed on the s o u t h e r l y s i d e s o f them. s i t e s v a r i e d w i t h snow depth.  S e l e c t i o n o f bed  As snow became deeper, moose  bedded c l o s e r t o l a r g e r t r e e s i n the denser c a n o p i e d p a r t s of f o r e s t stands. protected  Moose showed g r e a t e r s e l e c t i o n f o r  s i t e s as w i n t e r c o n d i t i o n s became more  severe.  Secondary s e r a i s u c c e s s i o n was examined w i t h r e s p e c t t o s e v e r a l a t t r i b u t e s f o r mesic environments on the two commonest s u b s t r a t e s , g l a c i a l t i l l  and l a c u s t r i n e d e p o s i t s .  iv F l o r i s t i c s of s e r a i stages from 1-200 l a c u s t r i n e s o i l s , v e g e t a t i o n was deciduous  phase was  prolonged.  y e a r s r e v e a l e d t h a t on  more, d i v e r s e and  Species d i v e r s i t y d e c l i n e d  around y e a r 25 on t i l l b u t n o t on l a c u s t r i n e . changes o c c u r r e d i n the t r e e l a y e r : l a y e r developed  S e v e r a l major  f i r s t , a deciduous  e s p e c i a l l y on l a c u s t r i n e s o i l s ;  a f t e r 25 y e a r s on t i l l  the  (45 y e a r s o r more on  tree  second,  lacustrine),  l o d g e p o l e p i n e became most abundant; t h i r d , p i n e  was  g r a d u a l l y r e p l a c e d by w h i t e spruce a f t e r 150-200 y e a r s o r more; f o u r t h , s u b a l p i n e f i r would p r o b a b l y become the dominant t r e e s p e c i e s i n the absence o f f i r e . phytomass, though c o n t r i b u t i n g l i t t l e  Understory  to the mature f o r e s t  mass, i n c r e a s e d d r a m a t i c a l l y . t o peaks e a r l y i n s u c c e s s i o n and then remained low.  Approximate n e t p r i m a r y p r o d u c t i o n  o f the u n d e r s t o r y on t i l l was g r e a t e s t a t age 11 w i t h 2 2 g/m / y r produced and l e a s t a t age 39 w i t h 18 g/m / y r produced.  133  U n d e r s t o r y p r o d u c t i o n i n the mature f o r e s t was  an  2  e s t i m a t e d 27 g/m  /yr.  The  shrubs  c o n t r i b u t e d 70 p e r c e n t ,  26 p e r c e n t , 44 p e r c e n t , and 26 p e r c e n t o f annual  production  at ages 1, 11, 39, and 195 y e a r s , p o s t - d i s t u r b a n c e . p r o t e i n and l i g n i n v a l u e s were determined ( e i g h t s h r u b s , one c o n i f e r , one  f o r 10 s p e c i e s  l i c h e n ) f o r an annual  Crude p r o t e i n averaged 7 p e r c e n t . a n d  Crude  l i g n i n , 9.8  cycle.  percent.  Crude p r o t e i n i n c r e a s e d a b r u p t l y from steady w i n t e r v a l u e s t o peaks o f 10-15  p e r c e n t i n J u n e - J u l y and then r e t u r n e d t o  low l e v e l s by October.  Leaf p r o t e i n was  higher than,  and  V  p r e d i c t a b l e from, stem l e v e l s .  Crude p r o t e i n v a r i e d by-  s p e c i e s , sometimes by s u b s t r a t e  and r a r e l y by h a b i t a t - t y p e ,  at l e a s t f o r t h e s p e c i e s  analyzed.  The l i c h e n , l u n g w o r t ,  r e t a i n e d a h i g h p r o t e i n v a l u e o f a p p r o x i m a t e l y 11 p e r c e n t throughout the year.  Lignin levels varied  seasonally,  though l e s s d r a m a t i c a l l y than crude p r o t e i n . a f f e c t e d by s p e c i e s , s u b s t r a t e  and age o f s e r a i s t a g e .  P r o t e i n l e v e l s were s i m i l a r t o those r e p o r t e d literature.  Factors  L e v e l s were  i n the  i n f l u e n c i n g crude p r o t e i n were  d i f f i c u l t to disentangle  due t o c o n f o u n d i n g .  was s t u d i e d w i t h r e s p e c t  t o d i f f e r e n c e s i n snow  between h a b i t a t s .  climate  features  Moose moved i n t o w i n t e r ranges b e f o r e  snow depths were l i m i t i n g . t r i g g e r migration.  Winter  T h i s i n d i c a t e s snow a c t s t o  On w i n t e r  ranges, moose a l s o moved i n t o  f o r e s t e d h a b i t a t s i n m i d - w i n t e r (January) when snow depths approached 80 cm. habitats.  Snow depths and d e n s i t i e s v a r i e d between  Snow c o v e r was more v a r i a b l e i n p a r t i a l l y  c u t o v e r s than i n the open o r f o r e s t e d s t a n d s .  logged  The c l i m a t e  of f o r e s t , ecotone and a d j a c e n t open a r e a s were documented. Compared t o a d j a c e n t open a r e a s , the f o r e s t had h i g h e r r e l a t i v e h u m i d i t y , l e s s w i n d , more moderate temperatures and a p p r o x i m a t e l y 50 p e r c e n t o f t h e snow depth. zone from open t o f o r e s t c l i m a t e s narrow, l e s s than 50 m.  The t r a n s i t i o n  appeared t o be r e l a t i v e l y  The r e l a t i o n s h i p between c a r r y i n g  c a p a c i t y , h a b i t a t s e l e c t i o n and home range are d i s c u s s e d with reference  t o moose and management o f t h e i r h a b i t a t .  Management recommendations and s u g g e s t i o n s f o r f u t u r e r e s e a r c h are p r o v i d e d .  TABLE OF CONTENTS Page ABSTRACT  i i  TABLE OF CONTENTS  v i i  LIST OF TABLES  xiii  LIST OF FIGURES AND ILLUSTRATIONS  xx  LIST OF APPENDICES  xxiv  ACKNOWLEDGEMENTS  xxix  1.  2.  INTRODUCTION  1  1.1  The Study  1  1.2  The Approach t o the Study  6  1.3  The Need f o r I n t e g r a t e d  8  1.4  A Land Use P e r s p e c t i v e .  11  1.4.1  General Introduction  11  1.4.2  Mining  15  1.4.3  Agriculture.  18  1.4.4  Forestry  23  1.4.5  Wildfire  29  Management  THE STUDY AREAS  34  2.1  Biophysical Setting  34  2.2  The P r i m a r y Study Areas  52  2.3  The Secondary Study Areas  68  2.4  Moose D i s t r i b u t i o n and abundance  70  vii  viii 3.  HABITAT USE AND  SELECTION  3.1  Introduction  3.2  Methods  73 •  3.2.1  The S y n o p t i c Survey  3.2.2  P e l l e t Group C o u n t i n g Methods  3.2.3 3.3  73  80  The A e r i a l Surveys  81  Results  84  3. 3.1  H a b i t a t Use  84  3.3.2  Habitat Selection  97  3.3.3  The Timing o f M i g r a t i o n and  Discussion 3.4.1  4.  77  f o r D e t a i l e d Survey.  Occupany P e r i o d s 3.4  77  The Importance o f H a b i t a t Variability.  99 102 102  FOOD HABITS  109  4.1  Introduction  109  4.2  Methods  110  4.3  4.2.1  Rumen A n a l y s i s  110  4.2.2  Trailing  117  4.2.3  The Browsed Stem Survey  118  Results  118  4.3.1  The Range of S p e c i e s Taken  118  4.3.2  The S e a s o n a l Trends  12 0  The E f f e c t o f H a b i t a t - T y p e on D i e t Discussion 4.4.1 Methodology  125 130 130  4.3.3  4.4  ix Page 4.4.2  V a r i a t i o n s i n the D i e t  4.4.3  Some Management I m p l i c a t i o n s  4.4.4 5.  6.  7.  131  of V a r i a t i o n s i n the D i e t  133  F u t u r e Research. . . .  135  THE DYNAMICS OF WINTER BROWSING  137  5.1  Introduction  137  5.2  Methods  138  5.3  Results  143  5.3.1  The I n c i d e n c e  o f Use  14 3  5.3.2  The Time o f Use  149  5.3.3  The L e v e l o f Use  152  BED SITE SELECTION BY MOOSE IN WINTER  15 7  6.1  Introduction  15 7  6.2  Methods  15 9  6.3  Results  162  6.4  Discussion  179  SECONDARY SUCCESSION IN SUB-BOREAL FORESTS  184  7.1  Introduction  184  7.2  Methods  18 8  7.2.1  Stratification  188  7.2.2  F i e l d Sampling P r o c e d u r e s  190  7.2.3  The P r e d i c t i o n o f Mass and  7.2.4 7.3  H e i g h t o f Woody P l a n t s  198  Date-Analysis  207  R e s u l t s f o r M e s i c Upland S i t e s  208  7.3.1  20 8  The Data Base  X  Page 7.3.2 7.3.3 7.3.4  F l o r i s t i c Changes i n S e r a i Succession  210  Temporal Dynamics o f the Tree L a y e r  237  Phytomass, H e i g h t and B a s a l Area o f the Shrub L a y e r i n S e r a i P l a n t Communities  246  7.3.4.1 7.3.4.2  7.3.5 7.3.6  For the combination of s p e c i e s  246  Trends i n phytomass and h e i g h t o f food species  251  Phytomass o f the Herb L a y e r i n S e r a i P l a n t Communities  254  Net P r i m a r y Understory  258  P r o d u c t i v i t y o f the  7.4  Results f o r Riparian S i t e s  264  7.5  Discussion  269  7.5.1 7.5.2  P r e d i c t i n g Successional Development. Trends i n P r o d u c t i o n of Cover and Food  8.  269 2 76  NUTRITIVE ASPECTS OF MOOSE FORAGES  2 85  8.1  Introduction  2 85  8.2  Methods  288  8.3  Crude P r o t e i n L e v e l s  291  8.4  Lignin Levels  30 8  8.5  Discussion  314  8.5.1  Crude P r o t e i n L e v e l s i n Moose Forages  314  xi Page 8.5.2 8.5.3  A s s e s s i n g the. N u t r i t i v e V a l u e s of Forages Factors A f f e c t i n g Nutrient Levels  9.  318  322  EFFECTS OF FORESTS ON. WINTER CLIMATE  333  9.1  Introduction  333  9.2  Methods  335  9.2.1 9.2.2  9.2.3 9. 3  335  Habitat-Types  336  Climate  340  o f the F o r e s t Edge  Results 9.3.1 9.3.2  9.3.3 9.4  The E s t i m a t i o n . o f M i g r a t i o n and W i n t e r Range Occupancy . Snow C h a r a c t e r i s t i c s o f  341 M i g r a t i o n and Snow A c c u m u l a t i o n of H a b i t a t - T y p e s Snow C h a r a c t e r i s t i c s o f  341  Habitat-Types  345  C l i m a t e o f t h e F o r e s t Edge  355  Discussion 9.4.1 9.4.2  The Role o f Snow Pack i n I n i t i a t i n g Migration The Role of C l i m a t e i n  365 365  D i f f e r e n t i a l Use Between H a b i t a t s . . . . 367 10.  DISCUSSION Habitat Relationships i n Moose Management 10.2 The E f f e c t s o f Timber Management  379  10.1  379  on Moose H a b i t a t .  39 4  10.2.1 F e l l i n g  397  10.2.2 S i t e P r e p a r a t i o n  402  xii Page 10.2.3  Stand E s t a b l i s h m e n t  405  10.2.4  Stand Tending .  408  10.2.5  Stand P r o t e c t i o n  412  10.2.6  General  Management  Considerations. 10.3  Overview and Recommendations  414 416  11.  LITERATURE CITED. .  42 7  12.  APPENDICES  460  VITA  LIST OF TABLES TABLE 1.1  1.2  1.3  Page P o p u l a t i o n Growth and F u t u r e P r o j e c t i o n s f o r P r i n c e George and the S u r r o u n d i n g District Major Events i n the. S e t t l e m e n t and Growth of P r i n c e George and the S u r r o u n d i n g Region  14  16  Number and A r e a of. Farms, and Numbers of C a t t l e f o r the P r o v i n c e and f o r the P r i n c e George R e g i o n , 1881-1971  20  Trends i n Logging Methods and A r e a Cut i n the P r i n c e George F o r e s t D i s t r i c t , 1950-1973 . . .  27  E s t i m a t e d Areas of Broad V e g e t a t i o n C l a s s e s i n F i v e Major Drainages i n N o r t h - C e n t r a l B r i t i s h Columbia (from W h i t f o r d and C r a i g 1918)  31  2.1  C l i m a t i c Parameters f o r the Study Area  40  2.2  Major S o i l A s s o c i a t i o n s f o r the Study A r e a and T h e i r R e l a t i o n s h i p t o P a r e n t M a t e r i a l s and M o i s t u r e Regimes  44  E s t i m a t e d R e l a t i v e Abundance and Herd S t r u c t u r e f o r W i n t e r i n g Moose on the I n t e n s i v e Study Areas,, 1964-65 t o 1975-76  58  2.4  Types o f A n a l y s e s Conducted on the Study Area.. . .  69  3.1  R e s u l t s from T r i a l P e l l e t Group Survey: T i m e / P l o t and Number of G r o u p / P l o t  78  R e l a t i v e W i n t e r Use o f A v a i l a b l e H a b i t a t s on S e l e c t e d Study A r e a s , Based on P e l l e t Group Surveys  87  R e l a t i v e W i n t e r Use o f Major H a b i t a t - T y p e s i n the S u b - B o r e a l F o r e s t , Based on P e l l e t Group Surveys  89  1.4 1.5  2.3  3.2  3.3  xiii  xiv TABLE 3.4  3.5  3.6  3.7  3.8  Page W i n t e r Use o f Ecotones Between F o r e s t s and V a r i o u s l y Aged S e r a i S t a g e s , Based on P e l l e t Group Surveys W i n t e r U t i l i z a t i o n o f Roads and H a b i t a t s i n which they were L o c a t e d , Based on Accumulated P e l l e t Groups i n 1973 a t McKenzie.  92  . . 95  R e l a t i v e Summer Use o f . H a b i t a t - T y p e s on Accumulated Summer Feces Recorded i n t h e 1973 S y n o p t i c Survey. . .  96  D i s t r i b u t i o n o f P e l l e t Group P l o t s A c c o r d i n g t o H a b i t a t - T y p e and t h e Number o f Groups they C o n t a i n e d on t h e I n t e n s i v e Salmon Area  98  S e l e c t i o n o f H a b i t a t - T y p e s by Moose i n W i n t e r as I n d i c a t e d by Accumulated P e l l e t Groups on the I n t e n s i v e Salmon A r e a . .  98  4.1  Components o f Rumen O l i g e s t a A f t e r Sample Preparation  112  4.2  The E f f e c t o f A n a l y t i c a l Method on Frequency o f Occurrence o f P l a n t Taxa Recorded i n Moose Rumen Samples  114  4.3  The E f f e c t o f A n a l y t i c a l Method on Amounts o f P l a n t Taxa I d e n t i f i e d i n Moose Rumen Samples . . . 116  4.4  V a r i e t y o f P l a n t S p e c i e s Eaten by Moose, by Forage C l a s s , i n V a r i o u s P a r t s o f T h e i r N o r t h American Range . . .  119  Food H a b i t s o f Moose, i n N o r t h - C e n t r a l B r i t i s h Columbia, Based on T r a i l i n g and Rumen A n a l y s i s , 1971^-74 (%-basis)  122  4.5  4.6  Comparisons o f Food H a b i t s o f Moose Between D i f f e r e n t H a b i t a t s i n E a r l y and L a t e W i n t e r . . . . 126  4.7  W i n t e r Food P r e f e r e n c e s o f Moose i n N o r t h C e n t r a l B r i t i s h Columbia, by ^Habitat-Type  i 127  5.1  P r o p o r t i o n s o f Twigs t h a t were Browsed Once and Twice i n Major H a b i t a t s on the E a g l e , Grove and Salmon W i n t e r Ranges D u r i n g the 1972-73 W i n t e r  144  XV  TABLE 5.2  Page Number of Times P l a n t s of S u b a l p i n e F i r , Paper B i r c h , R e d - o s i e r Dogwood and W i l l o w were Browsed on the E a g l e , Grove and Salmon Study Areas d u r i n g the 1972-73 W i n t e r .  146  5.3  P r o p o r t i o n of P l a n t s Browsed.by S p e c i e s H a b i t a t , i n the 1972-73 W i n t e r  148  5.4  Time of Browsing and L e v e l of U t i l i z a t i o n (Weight-Basis) f o r A l l S p e c i e s , H a b i t a t Types, Study Areas and Months  6.1  and  15 0  Major H a b i t a t s , Snow Depth C l a s s e s and Study A r e a s Sampled f o r Bed S i t e E x a m i n a t i o n s  160  6.2  Example of Data Sheets-Used to Study Bed S i t e s . .  161  6.3  Time Spent by Moose i n Beds as I n d i c a t e d by Feces and U r i n e , A c c o r d i n g t o H a b i t a t and Month.  165  L o c a t i o n s of Moose.Beds w i t h Respect t o P o s i t i o n on S l o p e , and A s p e c t  166  Comparison of C o n i f e r S p e c i e s A v a i l a b l e as S h e l t e r T r e e s , W i t h Those -Used by Moose  169  O r i e n t a t i o n of Moose i n T h e i r Beds, and i n R e l a t i o n t o the S h e l t e r Tree  171  L o c a t i o n of Beds i n Quamaniqs, as A f f e c t e d by H a b i t a t and Snow Depth C l a s s  178  Comparison of Snow Depths between Moose Bedding S i t e s and A d j a c e n t Areas  179  S c a l e Used t o A s s e s s Canopy-Coverage of Understory Vegetation (Clayer) ( a f t e r Daubenmire 19 59), p l u s Domin S c a l e Equivalents  195  Summary of F e a t u r e s Sampled i n the S y n o p t i c Study o f S u c c e s s i o n  197  The E f f e c t o f S i t e on P r e d i c t i n g Mass f o r S e l e c t e d Shrub S p e c i e s . . •  201  6.4 6.5 6.6 6.7 6.8 7.1  7.2 7.3  xvi TABLE 7.4 7.5 7.6  7.7 7.8 7.9  7.10  7.11  7.12  7.13  Page 2 C o e f f i c i e n t s of D e t e r m i n a t i o n ( r v a l u e ) f o r S i x Independent V a r i a b l e s Used t o P r e d i c t Phytomass o f 19 S u b - B o r e a l Shrubs . . . . Regression C o e f f i c i e n t s f o r P r e d i c t i n g H e i g h t from Diameter Measurements  2 05  Regression C o e f f i c i e n t s f o r P r e d i c i n g Oven-Dried, Above-Ground Phytomass o f 19 S u b - B o r e a l Shrubs from Diameter, and from Diameter Squared by Length Measurements. A l l V a r i a b l e s Based.on L o g a r i t h m i c Transformed Data  206  D i s t r i b u t i o n o f Sampling S i t e s f o r the P l a n t S u c c e s s i o n Study. .  209  P l a n t Community Names f o r S u c c e s s i o n a l Stages on T i l l and L a c u s t r i n e S u b s t r a t e s  211  203  P e r c e n t Canopy-Coverage/Frequency o f Occurrence V a l u e s f o r Major P l a n t S p e c i e s of the Herb (C) L a y e r i n a S u b - B o r e a l F o r e s t Sere i n a M e s i c Environment on the T i l l Substrate. .  . 213  P e r c e n t Canopy-Coverage/Frequency o f Occurrence V a l u e s f o r Major P l a n t S p e c i e s of the Herb (C) L a y e r i n a S u b - B o r e a l F o r e s t Sere i n a M e s i c Environment on the Lacustrine Substrate. .  215  P e r c e n t Canopy-Coverage/Frequency o f Occurrence V a l u e s f o r Major P l a n t S p e c i e s of the Herb (C) L a y e r i n P a r t i a l l y Logged S u b - B o r e a l F o r e s t Stands i n a M e s i c Environment on T i l l and L a c u s t r i n e S u b s t r a t e s .  .217  P e r c e n t S p e c i e s C o m p o s i t i o n (Stem-Basis) o f the Shrub (B) L a y e r i n a S u b - B o r e a l F o r e s t Sere i n a M e s i c Environment Over T i l l Substrates  218  P e r c e n t S p e c i e s C o m p o s i t i o n (Stem-Basis) of the Shrub (B) L a y e r i n a S u b - B o r e a l F o r e s t Sere i n a M e s i c Environment Over L a c u s t r i n e Substrates  219  xvii x  Page  P e r c e n t S p e c i e s C o m p o s i t i o n (Stem-Basis) of the Shrub (B) L a y e r i n P a r t i a l l y Logged Stands o f S u b - B o r e a l F o r e s t s i n a M e s i c Environment  220  Temporal Changes i n Tree S p e c i e s Composition f o r Mesic Sub-Boreal F o r e s t s on T i l l and L a c u s t r i n e S u b s t r a t e s  238  Temporal Trends f o r C o n i f e r o u s R e g e n e r a t i o n i n a M e s i c Environment Over T i l l and Lacustrine Substrates  240  Temporal Changes i n C o m p o s i t i o n and P r o p o r t i o n s o f Dead Trees i n M e s i c SubB o r e a l F o r e s t Stands on T i l l and L a c u s t r i n e Substrates  242  Temporal Changes i n B a s a l A r e a , Canopy C l o s u r e , and H e i g h t of Dominant Trees i n M e s i c S u b - B o r e a l F o r e s t s on T i l l and Lacustrine Substrates  244  S t a t i s t i c s (mean = sd) f o r the Shrub (B) L a y e r i n M e s i c S u b - B o r e a l Seres Over T i l l and L a c u s t r i n e S u b s t r a t e s  247  Trends i n Phytomass o f the Herb (C) L a y e r f o r Seres i n M e s i c S u b - B o r e a l F o r e s t s on T i l l and L a c u s t r i n e S u b s t r a t e s  256  TABLE 7.14  7.15  7.16  7.17  7.18  7.19  7.2 0  7.21  Approximate Net P r i m a r y P r o d u c t i v i t y o f U n d e r s t o r y V e g e t a t i o n (Layers B and C) a t Four S u c c e s s i o n a l Stages on T i l l S u b s t r a t e . . . . 260  7.22  Changes i n P r o p o r t i o n s o f P l a n t Components of S e l e c t e d Shrub S p e c i e s a t Four S u c c e s s i o n a l Stages on T i l l S u b s t r a t e s  264  Major F e a t u r e s of S e r a i Stages i n F o r e s t S u c c e s s i o n on R i p a r i a n ( A l l u v i a l ) H a b i t a t s (Adapted from Sumanik (1968) and Waring (1970) )  267  L o c a t i o n , H a b i t a t , S u b s t r a t e , and S p e c i e s C o l l e c t e d f o r Crude P r o t e i n and L i g n i n Analyses  2 89  7.2 3  8.1  xviii TABLE 8.2 8.3  8.4  8.5  8.6  8.7  8.8 8.9  8.10  9.1 9.2  9.3 9.4  Page Estimates of Experimental Error i n Protein Analyses f o r S e l e c t e d Species..  291  Crude P r o t e i n and L i g n i n L e v e l s i n Major Moose Forages Averaged Over an Annual C y c l e (May 1972 - A p r i l 1973)  292  Crude P r o t e i n and L i g n i n L e v e l s i n the C u r r e n t Year's Stems and Leaves of S e l e c t e d Browse S p e c i e s  295  P r o p o r t i o n s o f Stem and L e a f T i s s u e i n C u r r e n t Annual Growth o f S e l e c t e d Browse S p e c i e s C o l l e c t e d i n September 19 72  297  Comparison o f P r o t e i n and L i g n i n Contents o f W i l l o w and Paper B i r c h C o l l e c t e d from T i l l and L a c u s t r i n e S u b s t r a t e s of the Burn H a b i t a t a t the Grove Study A r e a  299  E f f e c t of H a b i t a t on Crude P r o t e i n L e v e l s i n S e l e c t e d Browse S p e c i e s a t the E a g l e and Salmon Study Areas . . . . .  303  Y e a r - t o - Y e a r V a r i a t i o n s i n the Content of Crude P r o t e i n and L i g n i n  307  D i f f e r e n c e i n P e r c e n t L i g n i n Content as A f f e c t e d by S u b s t r a t e , H a b i t a t and Stand Age  312  Comparison o f Crude P r o t e i n V a l u e s f o r C u r r e n t Annual Growth o f Common W i n t e r Foods o f Moose (November-March)  316  L o c a t i o n , S i t e Number, E l e v a t i o n , and H a b i t a t of Snow Courses. .  337  Annual V a r i a t i o n s i n Snow Depths i n Open o r Deciduous F o r e s t H a b i t a t s a t E a g l e , Grove and Salmon W i n t e r Ranges f o r March 1972, 1973, and 1974  346  Monthly Snow Depths and D e n s i t i e s f o r Three H a b i t a t s on the E a g l e W i n t e r Range  34 8  Monthly Snow Depths f o r Four H a b i t a t s on the Grove W i n t e r Range  349  xix TABLE 9.5 9.6 9.7 9.8  9.9  Page Monthly Snow D e n s i t i e s f o r Four H a b i t a t s on the Grove W i n t e r Range.  350  Monthly Snow Depths, f o r F i v e H a b i t a t s a t the Salmon W i n t e r Range.  351  Monthly Snow D e n s i t i e s . f o r F i v e H a b i t a t s a t the Salmon W i n t e r Range  352  Comparison o f S e l e c t e d C l i m a t i c Parameters Between the S o u t h - F a c i n g Ecotone a t t h e Grove Study A r e a ( S t a t i o n a t 76 m i n the Open) and P r i n c e George. A i r p o r t .  357  Mean Monthly Temperature, R e l a t i v e H u m i d i t y , Snow Pack, and Wind f o r C l e a r c u t and A d j a c e n t F o r e s t S i t e s a t t h e Bowron Study A r e a , 1972-73 Data  359  9.10 Mean Monthly Depth, D e n s i t y , and P e n e t r a n c e of Snow A c r o s s t h e S o u t h - F a c i n g , F o r e s t Burn Ecotone a t t h e Grove Study A r e a , 1972-73 W i n t e r 10.1 10.2  363  G e n e r a l F e a t u r e s o f B a s i c Resources R e q u i r e d by Moose  395  S o i l D i s t u r b a n c e and S l a s h A c c u m u l a t i o n s R e s u l t i n g from D i f f e r e n t Types o f L o g g i n g i n Western N o r t h America ( D e r i v e d from Bockheim e t a l . 1975)  400  LIST OF FIGURES FIGURE 1.1  1.2  1.3 2.1 2.2  2.3  2.4  2.5 2.6 2.7 2.8  Page The g e n e r a l r e l a t i o n s h i p of sub-models t h a t comprise a moose-forest model. D e r i v e d from Haagenrud and H j e l j o r d (1976) and Houston (1968) Development of f o r e s t r y i n the P r i n c e George F o r e s t D i s t r i c t as i n d i c a t e d by the annual c u t and the number of o p e r a t i n g s a w m i l l s , 1914-1974 2 Annual a r e a (km ) burned by w i l d f i r e i n the P r i n c e George F o r e s t D i s t r i c t , 1910-1975  4  25  32  L o c a t i o n s o f the study a r e a s , and o f p l a c e names mentioned i n the t e x t  35  Longterm monthly averages o f some temperature and p r e c i p i t a t i o n p a r a meters f o r the P r i n c e George weather station  38  O b l i q u e a e r i a l photographs i l l u s t r a t i n g the g e n e r a l t e r r a i n and v e g e t a t i o n o f the P r i n c e George study a r e a  43  A schematic i l l u s t r a t i o n o f the major s o i l a s s o c i a t i o n s i n the study a r e a , and t h e i r t o p o g r a p h i c r e l a t i o n s h i p t o each o t h e r  45  Photographs of the E a g l e , Grove and Salmon w i n t e r ranges  53  A s o i l a s s o c i a t i o n map study a r e a  55  A f o r e s t cover map area  of the E a g l e  o f the E a g l e  A s o i l a s s o c i a t i o n map study a r e a  study 56  of the Grove 61 xx  xx i FIGURE 2.9 2.10 2.11 3.1  3.2  3.3  3.4  3.5  4.1  5.1  6.1  6.2  Page A f o r e s t cover map o f t h e Grove study a r e a  63  A s o i l a s s o c i a t i o n map o f t h e Salmon study area  65  A f o r e s t cover map o f the Salmon study a r e a  67  Photographs i l l u s t r a t i n g logged h a b i t a t s i n sub-boreal f o r e s t s : a) s e l e c t i v e , b) c u t and l e a v e , and c) c l e a r c u t  76-  The r e l a t i o n s h i p between d a i l y s n o w f a l l and the t i m i n g o f the a e r i a l t r a n s e c t s u r v e y s , January 19 72 t o May 19 7 3  83  A map showing f l i g h t l i n e s used f o r t h e a e r i a l t r a n s e c t surveys on the Grove study a r e a  85  R e l a t i v e use by moose o f ecotones and a d j a c e n t h a b i t a t s , based on p e l l e t group t r a n s e c t s  93  Number o f moose seen/minute o f f l y i n g on the E a g l e , Grove and Salmon study areas d u r i n g a e r i a l t r a n s e c t surveys i n the 1972-73 w i n t e r  101  The s e a s o n a l changes i n f o r a g e c l a s s e s eaten by moose i n n o r t h - c e n t r a l B r i t i s h Columbia  124  Photographs i l l u s t r a t i n g t h e methods o f t a g g i n g t w i g s and measuring diameter a t p o i n t o f browsing  141  The r e l a t i o n s h i p between snow depth and the l e n g t h o f time moose spent i n beds, as i n d i c a t e d by r e l a t i v e amounts o f f e c e s and u r i n e  163  The r e l a t i o n s h i p between snow depth and the d i s t a n c e between bed s i t e s and t h e i r associated shelter trees  174  xxii FIGURE 6.3  6.4 7.1 7.2  7.3  7.4  7.5  7.6  7.7  7.8  8.1  Page The r e l a t i o n s h i p between snow depth and the d i f f e r e n c e i n crown c l o s u r e between a bed s i t e and the f o r e s t s t a n d i n w h i c h i t was l o c a t e d . . . .  176  An i l l u s t r a t i o n of e f f e c t i v e snow i n t e r c e p t i o n by the f o r e s t canopy  177  The s i t e and s t a t i o n l a y o u t used t o study secondary p l a n t s u c c e s s i o n  192  Photographs i l l u s t r a t i n g s e l e c t e d s u c c e s s i o n a l stages on mesic t i l l and lacustrine substrates  221  Some t r e n d s i n the f o r e s t s t a n d f e a t u r e s o f b a s a l a r e a , dominant t r e e h e i g h t , and canopy c l o s u r e i n s u b - b o r e a l f o r e s t s e r e s on t i l l and l a c u s t r i n e s u b s t r a t e s  245  Trends i n h e i g h t and mass o f browse and non-browse s p e c i e s i n s u b - b o r e a l f o r e s t s u c c e s s i o n on t i l l and l a c u s t r i n e substrates  252  P e r c e n t a g e c o m p o s i t i o n , by forage c l a s s , of phytomass i n the "C" o r Herb l a y e r a t your s u c c e s s i o n a l stages o f the subb o r e a l f o r e s t on t i l l and l a c u s t r i n e substrates  255  P e r c e n t a g e c o m p o s i t i o n , by forage c l a s s , o f the n e t p r i m a r y p r o d u c t i o n of the u n d e r s t o r y v e g e t a t i o n ( l a y e r s B and C) a t f o u r s u c c e s s i o n a l stages o f t h e subb o r e a l f o r e s t on the t i l l s u b s t r a t e  259  P e r c e n t a g e c o m p o s i t i o n , by s p e c i e s , o f the n e t p r i m a r y p r o d u c t i o n o f the shrubs ( l a y e r s B and C) a t f o u r s u c c e s s i o n a l stages o f the s u b - b o r e a l f o r e s t on the t i l l s u b s t r a t e  262  Photographs i l l u s t r a t i n g p l a n t s u c c e s s i o n a l stages on r i p a r i a n ( a l l u v i a l ) substrates  268  Crude p r o t e i n l e v e l s i n major p l a n t s p e c i e s eaten by moose i n s u b - b o r e a l forests  293  xxiii FIGURE 8.2  8.3  8.4  8.5 8.6  Page Comparisons o f crude p r o t e i n l e v e l s i n w i l l o w and s u b a l p i n e f i r growing on s i m i l a r s u b s t r a t e s but i n stands of d i f f e r e n t ages  301  Comparisons of crude p r o t e i n l e v e l s i n d i f f e r e n t s p e c i e s growing a t the same s i t e s  304  C o n s i s t e n c y i n crude p r o t e i n l e v e l s between aspen and w i l l o w f o r t h r e e different sites  306  L i g n i n l e v e l s i n major shrub s p e c i e s eaten by moose i n s u b - b o r e a l f o r e s t s  310  E f f e c t o f s i t e on l i g n i n l e v e l s i n s e l e c t e d s u b - b o r e a l shrubs  313  9.1  Photographs showing the use o f the w e s t e r n snow sampler and the penetrometer. . . . 338  9.2  P a t t e r n s o f snow a c c u m u l a t i o n and snow m e l t f o r the E a g l e , Grove and Salmon study a r e a s , and f o r weather s t a t i o n s a t P r i n c e George and A l e z a Lake  342  Some temperature and r e l a t i v e h u m i d i t y g r a d i e n t s a c r o s s the f o r e s t - o p e n ecotone a t the Grove study a r e a d u r i n g the 1972-73 w i n t e r  358  Wind run and snow depths a c r o s s the f o r e s t - o p e n ecotone a t the Grove study a r e a d u r i n g the 1972-73 w i n t e r  361  Major f a c t o r s and how they i n t e r - r e l a t e moose p o p u l a t i o n l e v e l s ( m o d i f i e d from Houston 1968)  384  9.3  9.4  10.1  LIST OF APPENDICES Page APPENDIX A. APPENDIX B.  APPENDIX C.  SCIENTIFIC AND COMMON NAMES OF PLANT SPECIES RECORDED IN THE STUDY AREA  4 60  SCIENTIFIC AND COMMON -NAMES OF BIRD AND MAMMAL SPECIES MENTIONED FOR THE STUDY AREA  467  STATISTICAL DATA USED FOR THE INTRODUCTION (SECTION 1)  469  , - Table C - l . E s t i m a t e d Annual Economic V a l u e o f M i n e r a l s P r o d u c e d . i n the Omineca M i n i n g D i s t r i c t , 1926-1974. . . . . 470 • . ••  Table C-2. Number and T o t a l A r e a o f Farms, and Number o f C a t t l e f o r the P r o v i n c e and f o r the P r i n c e George Region, 1881-1971  471  T a b l e C-3. Annual Cut o f Timber ( A l l S p e c i e s ) and the S a w m i l l s O p e r a t i n g i n the P r i n c e George F o r e s t D i s t r i c t , 1909-1975  472  Table C-4. A r e a o f F o r e s t Land D i s t u r b e d by W i l d f i r e s and by L o g g i n g i n the P r i n c e George F o r e s t D i s t r i c t , 1912-1975  473  APPENDIX D. - HABITAT USE AND SELECTION DATA (SECTION 3)  474  Table D - l . I n d i v i d u a l P l o t Data on Time and Number o f Accumulated P e l l e t Groups Counted in- the A p r i l , 19 72 T r i a l Used t o Determine the P e l l e t Group Survey Method  4 75  Table D-2. Example o f the R e c o r d i n g Format Used f o r the A e r i a l T r a n s e c t s o f the I n t e n s i v e Study A r e a s , and the Type of Data Recorded  477  xxiv  XXV  Page Table D-3. Example o f the Summary D e r i v e d from A e r i a l T r a n s e c t Data .  479  Table D-4. Data from P e l l e t Group T r a n s e c t s f o r S y n o p t i c Surveys i n 19 72. Table D-5. Data from P e l l e t Group T r a n s e c t s f o r S y n o p t i c Surveys i n 1973. APPENDIX E.  CHARACTERISTICS OF SAMPLES COLLECTED FOR THE FOOD HABITS STUDY (SECTION 4) . Table E - l . Rumen Samples: Date o f K i l l , Sex, Age,- and L o c a t i o n o f K i l l . .  APPENDIX F.  SUCCESSION DATA (FOR SECTION 7) Table F - l . P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f Major* P l a n t S p e c i e s Recorded i n the Herb L a y e r a t t h e S u c c e s s i o n Study S i t e s , MF2-MF7. .  491  T a b l e F-2. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f Major* P l a n t S p e c i e s Recorded i n the Herb L a y e r a t the S u c c e s s i o n Study S i t e s , MF8-MF13 . . 494 Table F-3. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r at t h e S u c c e s s i o n Study S i t e s , MF14-MF19. . 497 Table F-4. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r at t h e S u c c e s s i o n Study S i t e s , MF20-MF22. . 500 Table F-5. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r a t the S u c c e s s i o n Study S i t e s , SR1-SR6. . . 503 T a b l e F-6. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r a t t h e S u c c e s s i o n Study S i t e s , SR7-SR12 . . 506  xxvi Page Table F-7. Percentage Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r at the S u c c e s s i o n Study S i t e s , SR13-SR18. . 509 Table F-8. P e r c e n t a g e Canopy-Coverage/ Frequency o f Occurrence V a l u e s o f M a j o r * P l a n t S p e c i e s Recorded i n the Herb L a y e r a t the S u c c e s s i o n Study S i t e s , SR19-SR23. . 512 2 Table F-9. Phytomass (g/m , Oven-Dried B a s i s ) o f the Shrub L a y e r , by S p e c i e s , i n a S u b - B o r e a l F o r e s t Sere i n a M e s i c Environment on T i l l S u b s t r a t e s 515 2 Table F-10. Phytomass (g/m , Oven-Dried B a s i s ) o f the Shrub L a y e r , by S p e c i e s , i n a S u b - B o r e a l F o r e s t Sere i n a Mesic Environment on L a c u s t r i n e S u b s t r a t e s . . . . 516 2 Table F - l l . Phytomass (g/m , Oven-Dried B a s i s ) o f the Shrub L a y e r , by S p e c i e s , i n P a r t i a l l y Logged Sub-Boreal F o r e s t s i n a Mesic Environment on T i l l and Lacustrine Substrates 517 2 2 Table F-12. B a s a l A r e a (cm /m ) o f the Shrub L a y e r , by S p e c i e s , i n a Sub-Boreal F o r e s t Sere i n a Mesic Environment on T i l l Substrates 518 2 2 Table F-13. B a s a l A r e a (cm /m ) of the Shrub L a y e r , by S p e c i e s , i n a SubB o r e a l F o r e s t Sere i n a Mesic Environment on L a c u s t r i n e S u b s t r a t e s 520 2 2 Table F-14. B a s a l A r e a (cm /m ) o f the Shrub L a y e r , by S p e c i e s , i n P a r t i a l l y Logged Sub-Boreal F o r e s t s i n a M e s i c Environment on T i l l and L a c u s t r i n e Substrates 521 Table F-15. H e i g h t (cm) o f the Shrub L a y e r , by S p e c i e s , i n a Sub-Boreal F o r e s t Sere i n a Mesic Environment on T i l l Substrates 522  xxvii Page  APPENDIX G.  APPENDIX H.  Table F-16. H e i g h t (cm) o f the Shrub L a y e r , by S p e c i e s , i n a Sub-Boreal F o r e s t Sere i n a M e s i c Environment on Lacustrine Substrates  524  T a b l e F-17. H e i g h t (cm) o f the Shrub L a y e r , by S p e c i e s , i n P a r t i a l l y Logged Sub-Boreal F o r e s t s i n a Mesic Environment on T i l l and L a c u s t r i n e Substrates  525  Table F-18. Number of Stems Sampled i n the Shrub L a y e r , by S p e c i e s , i n a Mesic S u b - B o r e a l F o r e s t Sere on T i l l Substrates  526  Table F-19. Number o f Stems Sampled i n the Shrub L a y e r , by S p e c i e s , i n a Mesic S u b - B o r e a l F o r e s t Sere on Lacustrine Substrates  527  Table F-20. Number of Stems Sampled i n the Shrub L a y e r , by S p e c i e s , i n P a r t i a l l y Logged, M e s i c Sub-Boreal F o r e s t s on T i l l and L a c u s t r i n e Substrates  528  Table F-21. Oven-Dried Weights o f Components o f Major Shrub S p e c i e s i n Sub-Boreal F o r e s t s  529  DATA FOR NUTRIENT CONTENTS OF SAMPLED PLANT SPECIES (SECTION 8) . . .  534  Table G - l . Crude P r o t e i n L e v e l s (%) i n P l a n t Samples C o l l e c t e d from the P r i n c e George Study A r e a , A p r i l 1972 to A p r i l 1973 . . . '  535  Table G-2. L i g n i n V a l u e s (%) i n P l a n t Samples C o l l e c t e d from the P r i n c e George Study A r e a , A p r i l 1972 to A p r i l 1973  541  CLIMATIC DATA USED FOR  545  SECTION 9  Table H - l . Penetrance V a l u e s (1-11 S c a l e ) f o r Snow Hardness E s t i m a t e s A c r o s s the S o u t h - F a c i n g Ecotone a t the Grove A r e a , 1973  546  xxviii Page  APPENDIX I .  Table H-2. Monthly Means f o r Temperature and R e l a t i v e H u m i d i t y A c r o s s the SouthF a c i n g F o r e s t - B u r n Ecotone a t the Grove Study A r e a , 1972-73 W i n t e r  547  Table H-3. Wind Run (km/day) A c r o s s the S o u t h - F a c i n g F o r e s t - B u r n Ecotone a t Grove Study A r e a and the Exposed Burn S i t e a t Buckhorn  548  Table H-4. Snow Depths (cm) A c r o s s the West-Facing Ecotone a t the Grove Study A r e a , 19,72-73 W i n t e r  548  GLOSSARY OF TERMS AND USED IN THE TEXT  549  ABBREVIATIONS  C  ACKNOWLE DGEMENTS I t i s a pleasure  t o acknowledge the many persons who  contributed to t h i s project.  Thanks go t o my committee,  Drs. P. J . Bandy, F. B u n n e l l , I . M. Cowan, V. C. R u n e c k l e s , J . H. G. Smith and K. Sumanik, f o r t h e i r a d v i c e and c r i t i c a l review  o f my work.  To my s u p e r v i s o r , Dr. V. C.  " B e r t " B r i n k , I e x p r e s s my g r a t i t u d e f o r h i s i n t e l l e c t , p a t i e n c e , p e r s p e c t i v e , diplomacy and honesty. e n r i c h i n g experience  t o be one o f h i s graduate  I t was an students.  My study was f i n a n c e d , encouraged and f a c i l i t a t e d l a r g e l y by t h e B.C. F i s h and W i l d l i f e Branch. W i l d l i f e personnel  F i s h and  i n the P r i n c e George r e g i o n s u p p o r t e d and  a s s i s t e d me i n many ways.  Ken Sumanik encouraged me t o study  the p r o b l e m , and p r o v i d e d me w i t h t h e b e n e f i t o f h i s experience  and t h e w i t o f h i s i n s i g h t .  mine o f i n f o r m a t i o n .  Conservation  M i l t Warren was a  O f f i c e r s D. A d o l p h ,  B. C l a p p , L. Cox, W. Richmond, D. T u r n e r , G. V i n c e n t i n t r o d u c e d me t o t h e i r d i s t r i c t s , c o l l e c t e d rumen samples, and a c t e d as e x c e l l e n t g u i d e s and s o u r c e s o f i n f o r m a t i o n . Thanks a l s o go t o P. Brade, K. C h i l d , K. F u j i n o and R. Goodlad o f the P r i n c e George o f f i c e , and t o many i n the V i c t o r i a o f f i c e f o r t h e i r v a r i o u s c o n t r i b u t i o n s t o my  study.  Members o f t h e B.C. F o r e s t S e r v i c e p r o v i d e d much xxix  XXX  u s e f u l i n f o r m a t i o n on f o r e s t r y r e l a t e d a f f a i r s . acknowledge  I wish to  C. P. Axhorn, J . B u l l e n , R. C l i f f o r d ,  D.  G i l b e r t , E. Lemon, J . R e v e l , W. Young and the r a n g e r s t a f f s at P r i n c e George, H i x o n , Summit Lake and A l e z a Lake. I r e c e i v e d v a l u a b l e h e l p from the Resource A n a l y s i s Branch o f the Environment M i n i s t r y Inventory).  ( f o r m e r l y the B.C.  Land  I n p a r t i c u l a r , I thank Greg Cheeseman, A l  Dawson, the l a t e A l L u c k u r s t , Gary Runka and J i m van Barneveld. F o r e s t companies  i n the a r e a g e n e r o u s l y p r o v i d e d  maps, i n f o r m a t i o n and a s s i s t a n c e .  I wish to  acknowledge  Rustad B r o t h e r s Lumber Company and t h e i r f o r e s t e r ,  Don  Frood; H o l g e r Thomsen; Northwood P u l p and Timber Company Ltd.;  P r i n c e George P u l p and Paper Company L t d . ; and  Weldwood o f Canada Company L t d . The f o l l o w i n g people a s s i s t e d me i n d a t a - c o l l e c t i o n , b o t h i n t e d i o u s l a b o r a t o r y a n a l y s e s and i n mosquito-plagued, patience-demanding f i e l d work:  R i c k Bonar, Dave Dunbar,  C h r i s Easthope, O l l i e F r i c k e , John K e l l y , Ben Koop, M a r g a r e t L a r k i n , Mike Masson, W i l l a Noble, Sharon  Russell,  E r i c R u t t , Rankin S m i t h , Don Stevenson and C h r i s Whyte. John K e l l y d e s e r v e s s p e c i a l thanks b o t h f o r h i s i n v a l u a b l e h e l p and h i s f r i e n d s h i p . Other o r g a n i z a t i o n s who p r o v i d e d h e l p : of New  The  College  C a l e d o n i a and the Canada A g r i c u l t u r e E x p e r i m e n t a l  Farm f o r l a b o r a t o r y space and f a c i l i t i e s ;  the Canadian  xxxi W i l d l i f e Service,, f o r s c h o l a r s h i p s . Other i n d i v i d u a l s I w i s h t o acknowledge a r e : Bower, master o f the Cessna 185; R i c h a r d R e v e l , who d e s c r i b e d the s u b - b o r e a l spruce zone t o me;  Les first  Ed T e l f e r , whose  e x p e r i e n c e , common sense and i d e a s o f f e r e d example and s t i m u l a t i o n ; John P o w e l l and Douglas G o l d i n g o f the Canada F o r e s t r y S e r v i c e , f o r t h e i r d a t a and a d v i c e on  forest  c l i m a t o l o g y ; J i m Peek, f o r a d v i c e on snow measurements and s e v e r a l a s p e c t s of moose e c o l o g y ; Rod S i l v e r , f o r u s e f u l d i s c u s s i o n s o f what makes moose " t i c k " ; R i c k E l l i s , f o r i n f o r m a t i o n and d i s c u s s i o n on p l a n t s u c c e s s i o n ; and  Ralph  R i t c e y , f o r a d v i c e and comment on my i d e a s and w r i t i n g . For making the f i n a l  copy o f the t h e s i s so r e a d a b l e  thanks go t o B a r b a r a Smith, who  d i d an e x c e l l e n t job of  t y p i n g ; and t o L a u r a F r i i s , who  competently p r e p a r e d  graphs and  the  figures.  As i n most f i e l d p r o j e c t s , many i n d i v i d u a l s  and  f a m i l i e s p r o v i d e d a human environment which complemented and enhanced my e x p e r i e n c e s w h i l e we  l i v e d i n P r i n c e George.  w i s h t o acknowledge i n p a r t i c u l a r the f o l l o w i n g f a m i l i e s  I and  i n d i v i d u a l s : .the C l a p p s , F r o o d s , Gagnons, J a r o s c h s , Manns, P a g e t s , John S a w i t s k y , S p u r r s and Sumaniks. L a s t , b u t c e r t a i n l y n o t l e a s t , I w i s h t o thank family:  my  the s u p p o r t and f o r e b e a r a n c e of my w i f e , E l a i n e ,  and my two c h i l d r e n , Jenny and S t u a r t ; i n many r e s p e c t s t h i s t h e s i s i s as much t h e i r s as i t i s mine.  To my p a r e n t s , Ben  and B e r n i c e , I owe an unpayable debt.  My i n - l a w s , Len and  Mabel Weston, gave f r e e l y t h e i r u n d e r s t a n d i n g and s u p p o r t .  1.  INTRODUCTION  1.1  The  Study  Moose are e l u s i v e , s o l i t a r y u n g u l a t e s o f the b o r e a l f o r e s t s i n n o r t h - c e n t r a l B r i t i s h Columbia.  sub-  They have  been h i g h l y s u c c e s s f u l i n t h i s c o m p a r a t i v e l y h a r s h environment.  T h e i r success i s p r o b a b l y due t o t h r e e main  characteristics.  F i r s t l y , moose are adapted  to winters that  are l o n g , c o l d and snowy ( K e l s a l l 1969, K e l s a l l and 1971).  S e c o n d l y , they are browsers  Telfer  i n an a r e a where shrubs  and t r e e s form the major food r e s o u r c e , e s p e c i a l l y i n winter. species  F i n a l l y , they are a f i r e - o r s u c c e s s i o n a l l y - a d a p t e d ( G e i s t 1971)  and so can c a p i t a l i z e e f f e c t i v e l y  the superabundance o f f o r a g e produced f o r e s t s u c c e s s i o n ( T e l f e r 1974).  on  i n the e a r l y s t a g e s of  Since frequent f i r e s leave  l a r g e p o r t i o n s of b o r e a l and s u b - b o r e a l f o r e s t s i n these e a r l y stages  (Heinselman  1973  and o t h e r s ) , t h i s response i s  a d i s t i n c t advantage t o the s p e c i e s .  T h e i r response i s  m a n i f e s t e d by i n c r e a s i n g p r o d u c t i v i t y when n u t r i t i o u s abundant f o r a g e becomes a v a i l a b l e  ( G e i s t 1974,  and  Markgren  1969) . Moose are an i m p o r t a n t n a t u r a l r e s o u r c e . H i s t o r i c a l l y , i n the e a r l y 1900's, meat and h i d e s were used f o r food and c l o t h i n g by i n d i g e n o u s peoples and European 1  2 settlers.  S i n c e t h e n , the r e c r e a t i o n a l v a l u e o f moose has  i n c r e a s e d d r a m a t i c a l l y , e s p e c i a l l y i n the n o r t h - c e n t r a l region. (old  D u r i n g t h e 1970-74 p e r i o d , t h e P r i n c e George a r e a  management Areas 20-22) p r o v i d e d a p p r o x i m a t e l y an  average o f 16,000 man-days o f r e c r e a t i o n and a h a r v e s t o f 5,400 a n n u a l l y ( B r i t i s h Columbia F i s h and W i l d l i f e Branch 1970-1974).  Most r e c e n t l y , t h e non-consumptive use o f moose  has been r e c o g n i z e d as a growing  and i m p o r t a n t v a l u e .  A d d i t i o n a l l y , moose may w e l l p r o v i d e a p r o t e i n s o u r c e .  The  t e c h n i c a l f e a s i b i l i t y o f "game r a n c h i n g " moose has been demonstrated by Knorre  (197 4)  and o t h e r s .  Hence, moose w i l l  become an i n c r e a s i n g l y v a l u a b l e r e s o u r c e .  To d e a l  a d e q u a t e l y w i t h t h i s i m p o r t a n t r e s o u r c e , moose management must i n t e n s i f y . The  f o r e s t s t h a t moose c o l o n i z e d so s u c c e s s f u l l y  have a l t e r e d s i n c e moose f i r s t appeared.  T h i s change i s an  i n e v i t a b l e , n a t u r a l consequence o f f o r e s t s u c c e s s i o n .  The  s i g n i f i c a n t d i f f e r e n c e w i t h f u t u r e changes w i l l be the impact o f man and h i s a c t i v i t i e s .  Of these  pursuits,  h a r v e s t i n g t i m b e r and c o n t r o l l i n g f o r e s t f i r e s w i l l be t h e most i m p o r t a n t .  S i n c e these s u b - b o r e a l f o r e s t s a r e e a s i l y  a c c e s s i b l e and h i g h l y p r o d u c t i v e , f o r e s t - r e l a t e d development w i l l i n e v i t a b l y be w i d e l y d i s t r i b u t e d .  S i n c e moose a r e a l s o  w i d e l y spread i n these f o r e s t s , t h e q u e s t i o n a r i s e s , "What w i l l be t h e e f f e c t s o f human a c t i v i t i e s upon moose?" The purpose o f t h i s t h e s i s i s t o examine moose  3  habitat with p a r t i c u l a r reference related a c t i v i t i e s . their habitats  t o the e f f e c t s of f o r e s t -  Much i s known g e n e r a l l y about moose and  (Bedard e t a l . 1974).  However, l i t t l e i s  known about 1) moose h a b i t a t i n t h e n o r t h - c e n t r a l r e g i o n o f B r i t i s h Columbia, o r 2) t h e impact o f f o r e s t p r a c t i c e s upon this habitat.  These gaps a r e c r i t i c a l l i n k s i n a c h i e v i n g  i n t e g r a t e d management o f f o r e s t s and moose. C l e a r l y , a h a b i t a t - o r i e n t e d study does n o t d e a l a l l those components t h a t a f f e c t moose p o p u l a t i o n s .  with  Iti s  t h e r e f o r e u s e f u l t o p l a c e such a study i n c o n t e x t . V a r i a t i o n s i n moose p o p u l a t i o n s  a r e determined by two broad  t y p e s o f mechanisms (Houston 1968) - e n v i r o n m e n t a l and population.  The i n t e r f a c e o r l i n k a g e between them i s t h e  energy and n u t r i e n t s u p p l y moose.  ( i n c l u d i n g water) a v a i l a b l e t o  The a v a i l a b l e s u p p l y i s d e t e r m i n e d l a r g e l y by  e n v i r o n m e n t a l mechanisms, a l t h o u g h d e n s i t i e s and b e h a v i o u r of moose can o b v i o u s l y modify a v a i l a b i l i t y as w e l l .  The  f o c u s o f the p r e s e n t study i s on some o f t h e major e n v i r o n m e n t a l mechanisms t h a t determine t h e a v a i l a b l e s u p p l y of energy and n u t r i e n t s t o moose. schematically  i n Figure  T h i s i s shown  1.1.  The method by which I s t u d i e d these mechanisms i s r e f l e c t e d i n the o r g a n i z a t i o n of t h i s t h e s i s .  Four t o p i c s  make up t h e f i r s t p a r t o f the t h e s i s , v i z . , h a b i t a t use and s e l e c t i o n , d i e t , l e v e l o f use and bedding b e h a v i o u r .  In the  u l t i m a t e sense, t h e a v a i l a b i l i t y o f energy and n u t r i e n t s t o  3a  F i g u r e 1.1  The g e n e r a l r e l a t i o n s h i p of  sub-models  t h a t comprise a m o o s e - f o r e s t model. D e r i v e d from Haagenrud and (1976) and Houston  (1968).  Hjeljord  4  WILDLIFE MANAGEMENT DECISIONS  POPULATION MODEL  YEARLY HARVEST  CHANGES IN NATALITY, MORTALITY AND BODY WEIGHT.  BEHAVIOUR MODEL  hACTIVITY  HERD STRUCTURE, DENSITY AND NUMBER OF ANIMALS ENERGY MODEL  7  SATISFACTION GRAZING HABIT  COVER  FOREST/PATTERN/ PLANT SUCCESSION MODEL  FOOD ["AVAILABLE  .GRAZING QUANTITY 5T FOREST MANAGEMENT DECISIONS  •  TIMBER HARVEST  FOOD REQUIREMENT  GRAZING/ BROWSING MODEL  5 moose i s s e t by the c o n v e r s i o n o f s o l a r energy c h e m i c a l energy  stored i n plants.  i n t o the  However, n o t a l l o f t h e  v e g e t a t i o n i s f o r a g e f o r moose, and n o t a l l t h e f o r a g e i s a v a i l a b l e t o them.  Moreover, v e g e t a t i o n a l s o p r o v i d e s  escape c o v e r and s h e l t e r from t h e elements.  Thus the f i r s t  t a s k was an attempt t o p e r c e i v e the s u b - b o r e a l environment i n terms m e a n i n g f u l t o moose. h a b i t a t s were d e f i n e d by examining vegetation types. studies.  forest  Therefore,  t h e use and s e l e c t i o n of  Moose f o r a g e was a s s e s s e d by food h a b i t  Browse surveys were used t o e s t i m a t e t h e  p r o p o r t i o n o f t h e f o r a g e t h a t was e a t e n .  F i n a l l y , s i n c e the  a v a i l a b i l i t y o f s h e l t e r a f f e c t s t h e a v a i l a b i l i t y o f food ( B u n n e l l 1974), I a l s o examined t h e s e l e c t i o n o f bed s i t e s . These f o u r components make up t h e f i r s t p a r t o f t h e t h e s i s . Once t h e s e b i o l o g i c a l f e a t u r e s were d e f i n e d , I examined t h r e e major f a c t o r s t h a t m o d i f i e d them. t h r e e t o p i c s comprise  These  t h e second p a r t of t h e t h e s i s , v i z . ,  f o r e s t s u c c e s s i o n , f o r a g e n u t r i t i v e v a l u e s , and c l i m a t e . S u c c e s s i o n was s t u d i e d s i n c e i t determines  l o n g term  i n f o r a g e p r o d u c t i o n and t h e p r o v i s i o n o f c o v e r .  trends  Nutritive  a s p e c t s o f moose f o r a g e s were q u a n t i f i e d t o determine t h e r e l a t i o n s between s p e c i e s eaten and n u t r i e n t c o n t e n t , and t o a s s e s s what f a c t o r s i n f l u e n c e d n u t r i e n t l e v e l s .  C l i m a t e was  examined, e s p e c i a l l y snow, as i t i n f l u e n c e s the occupancy o f w i n t e r ranges, the a v a i l a b i l i t y o f f o r a g e , and the s e l e c t i o n and use o f h a b i t a t s .  These t h r e e t o p i c s t h a t comprise the  6 second p a r t o f t h e t h e s i s a r e a l s o m o d i f i e d  by f o r e s t  practices. The  t h i r d part of the t h e s i s , the D i s c u s s i o n ,  a t t e m p t s t o combine t h e f i r s t two p a r t s , t o a s s e s s t h e e f f e c t s o f f o r e s t r y a c t i v i t i e s on moose, and t o p r o v i d e an o v e r v i e w and recommendations f o r w i l d l i f e managers. 1.2 The  The Approach t o t h e Study  p r e s e n t management o f h a r v e s t a b l e  B r i t i s h Columbia has emphasized p o p u l a t i o n than h a b i t a t r e l a t i o n s h i p s . managers r e g u l a t e d  wildlife in  dynamics r a t h e r  U n t i l quite recently, w i l d l i f e  harvest through manipulation of hunting  seasons and bag l i m i t s .  Data were c o l l e c t e d from game and  hunter c h e c k s , h a r v e s t and h u n t e r q u e s t i o n n a i r e carry-over  and p o s t - s e a s o n c o u n t s .  to modify the production manipulation, recognized,  s u r v e y s , and  L i t t l e attempt was made  o f w i l d l i f e through h a b i t a t  a l t h o u g h t h e importance o f h a b i t a t was w e l l e.g., Smith  (1955).  In t h e 1960's, t h e t a c i t r e c o g n i t i o n o f h a b i t a t ' s r o l e e v o l v e d t o an a c t i v e p u r s u i t o f h a b i t a t management. Efficient  f o r e s t f i r e c o n t r o l and expanded l o g g i n g were t h e  two prime f a c t o r s i n d u c i n g  t h e development o f moose h a b i t a t  management programs i n n o r t h - c e n t r a l B r i t i s h Columbia.  The  i n c r e a s i n g e f f i c i e n c y o f p r o t e c t i n g s u b - b o r e a l f o r e s t s from wildfire vitiated successional  t h e main agent f o r c r e a t i n g e a r l y  ranges.  R e a l i z i n g t h e r e l a t i o n s h i p between  7 h i g h moose d e n s i t i e s and e a r l y s u c c e s s i o n a l s t a g e s , t h e adverse impact o f f i r e c o n t r o l on moose h a b i t a t s became r e a d i l y apparent.  The r a p i d l y i n c r e a s i n g acreage o f f o r e s t s  t h a t were logged was a l s o apparent.  Thus, t h e p r i s t i n e ,  u n c o n t r o l l e d agent o f range c r e a t i o n was p r o g r e s s i v e l y being  suppressed and r e p l a c e d by an agent whose impact was  c o n t r o l l e d by human d e s i g n .  Although creation or  m o d i f i c a t i o n o f moose h a b i t a t was a n c i l l a r y t o f o r e s t r y , b i o l o g i s t s r e a l i z e d t h a t h a b i t a t management f o r moose was readily feasible.  A passive regard  f o r vegetational  changes was t r a n s f o r m e d t o an a c t i v e i n t e r e s t i n m o d i f y i n g f o r e s t s by l o g g i n g f o r moose The  production.  broadened p e r s p e c t i v e o f w i l d l i f e management  from p r i m a r i l y a n i m a l s - o n l y a major development.  to animals-and-their-habitat  was  The growing awareness o f o p p o r t u n i t i e s  f o r h a b i t a t management were accompanied by t h e r e a l i z a t i o n t h a t moose h a b i t a t r e l a t i o n s h i p s i n s u b - b o r e a l f o r e s t s were l a r g e l y unknown, e x c e p t i n t h e most g e n e r a l  fashion.  E x i s t i n g i n f o r m a t i o n w i t h r e s p e c t t o logged h a b i t a t s was inadequate o r p o o r l y u n d e r s t o o d . P r e v i o u s moose s t u d i e s i n B r i t i s h Columbia were n e i t h e r w i t h i n the sub-boreal f o r e s t s nor d e a l t w i t h a s p e c t s o f moose e c o l o g y  other  ( H a t t e r 1950, Cowan e t a l . 1950,  Baynes 1956, R i t c e y and Verbeek 1969, Finnegan 1973). A l t h o u g h they p r o v i d e d  u s e f u l background i n f o r m a t i o n , t h e  major problem o f h a b i t a t r e l a t i o n s h i p s i n n o r t h - c e n t r a l  8 f o r e s t s was n o t addressed. Previous circumboreal  s t u d i e s o f moose h a b i t a t s elsewhere i n  ranges were a l s o u s e f u l as background  information  (e.g., Bergerud and Manuel 1968,  Lykke 1964,  Peek 1971,  Stevens 1970,  Peterson  T e l f e r 1967).  Houston 1968,  1 9 5 5 , , P i m l o t t 1961,  However, d e s p i t e r a t h e r  e x t e n s i v e s t u d i e s o f f o r e s t e d h a b i t a t s , logged areas have received s u r p r i s i n g l y l i t t l e attention.  Most s t u d i e s have  d e a l t w i t h n a t u r a l a r e a s , o r , where l o g g i n g had o c c u r r e d , i t r e c e i v e d almost i n c i d e n t a l a t t e n t i o n (e.g., Stevens 1970). Even i n s t u d i e s c a r r i e d o u t p r i m a r i l y on c u t o v e r s , e.g., Bergerud and Manuel (1968),  little  i n f o r m a t i o n on r e l a t i v e  use o f d i f f e r e n t t y p e s and ages o f c u t o v e r s has been provided.  S t u d i e s such as those r e c e n t l y p u b l i s h e d by Peek  e t a l . (1976)are e x c e p t i o n s 1.3  t o the r u l e .  The Need f o r I n t e g r a t e d Management  U n t i l q u i t e r e c e n t l y , s u p p l i e s o f t h e common property resources  exceeded demands.  Low p r i c e s f o r l a n d ,  g r a z i n g a l l o t m e n t s , c u t t i n g r i g h t s , and h u n t i n g l i c e n c e s , a l l r e f l e c t e d t h e apparent superabundance o f r e s o u r c e s .  In  f a c t , c o n s i d e r a b l e e f f o r t was expended by government t o encourage s e t t l e m e n t  and development.  Two c o n c o m i t a n t p r e s s u r e s situation.  radically altered this  Human p o p u l a t i o n i n c r e a s e d a t an e x p o n e n t i a l  r a t e , i n c r e a s i n g demands f o r goods and s e r v i c e s which i n t u r n  9 a c c e l e r a t e d demands f o r n a t u r a l r e s o u r c e s .  Many c i t i z e n s  e x p e r i e n c e d an i n c r e a s i n g amount of l e i s u r e time r e s u l t i n g from w e a l t h a c c r u e d through n a t u r a l r e s o u r c e developments i n the  region. These growing demands f o r a v a r i e t y of goods  s e r v i c e s , confronted  one  basic l i m i t :  a f i n i t e l a n d base.  As demands approached the l i m i t s imposed by  biophysical  p a r a m e t e r s , t h e y i n e v i t a b l y l e d t o c o n f l i c t and The  i n t e r e s t i n m u l t i p l e and  and  competition.  i n t e g r a t e d use was  an attempt  t o r e s o l v e t h e s e c o n f l i c t s i n a r a t i o n a l manner.  Whether  t h i s attempt w i l l be s u c c e s s f u l o r not i s s t i l l unknown. I t can be assumed t h a t an i n t e g r a t e d system of n a t u r a l r e s o u r c e a l l o c a t i o n and management i s a prime o p t i o n a v a i l a b l e t o the p u b l i c .  However, i n the p r e s e n t system,  i m p o r t a n t demands are u s u a l l y made by the p r o d u c e r s u s e r s of these r e s o u r c e s . s t a t e d as:  In s i m p l i s t i c terms, these can  defining objectives, devising a  process, inventorying of resources, f o l l o w e d by e v a l u a t i o n and  i m p l e m e n t a t i o n of p l a n s  re-adjustment.  terms so t h a t they can be e v a l u a t e d ,  The  objectives  integrated  E f f e c t i v e i n t e g r a t e d f o r e s t and  and time  and  wildlife  management must be dynamic r a t h e r than s t a t i c .  obvious.  of  i n operational  i n i t i a t e d w i t h i n the l i m i t s of a v a i l a b l e r e s o u r c e s ,  c o n t i n u a l re-assessment and  be  planning  i d e n t i f i e d u s e r groups must a l s o be d e f i n e d  technology.  and  The  need f o r  e v a l u a t i o n of programs i s  10 Competing demands f o r l i m i t e d r e s o u r c e s and t h e i r i n t e g r a t i o n has i m p o r t a n t e c o l o g i c a l i m p l i c a t i o n s .  First,  the o p t i o n o f " l e t t i n g n a t u r e take i t s c o u r s e " may appear l e s s t e n a b l e because n a t u r a l c a t a s t r o p h e s such as w i l d f i r e and i n s e c t o u t b r e a k s may s e r i o u s l y d i s r u p t t h e p r o d u c t i o n o f d e s i r e d and needed f o r e s t p r o d u c t s .  The apparent  impact o f  n a t u r a l c a t a s t r o p h e s on the economy may be d i r e c t l y to the degree o f r e s o u r c e commitment.  related  That s o c i e t y has  accepted t h i s r e l a t i o n s h i p i s c l e a r l y demonstrated by commitments t o f o r e s t f i r e s u p p r e s s i o n and i n s e c t c o n t r o l programs.  However, i t i s w o r t h n o t i n g t h a t economic  f a c t o r s may be as c a t a s t r o p h i c as n a t u r a l ones (K. Sumanik, p e r s . comm.). A second e c o l o g i c a l i m p l i c a t i o n i s t h a t human b e i n g s impose n o v e l means o f r e - d i s t r i b u t i n g and u s i n g r e s o u r c e s by a l t e r i n g n a t u r a l p a t t e r n s and p r o c e s s e s .  In  f o r e s t r y , wood p r o d u c t s w i t h t h e i r n u t r i e n t and energy c o n t e n t a r e m e c h a n i c a l l y removed and i n j e c t e d i n t o systems o f t e n f a r removed from t h e i r s o u r c e .  eco-  In w i l d l i f e  and f i s h e r i e s management, v e r t e b r a t e s p e c i e s a r e e x t r a c t e d by u n n a t u r a l methods and t r a n s p o r t e d t o o t h e r systems. None o f these i m p l i c a t i o n s i s n e c e s s a r i l y d e l e t e r i o u s t o t h e p r o d u c i n g systems, a l t h o u g h some l o g g i n g methods may cause n u t r i e n t d e p l e t i o n on poor s i t e s .  Since  we have g r e a t e x p e c t a t i o n s from n a t u r a l systems, and s i n c e we l i m i t o r a t l e a s t modify t h e i r f u n c t i o n i n g , i t i s  11 o b v i o u s t h a t we  s h o u l d u n d e r s t a n d how  manage them b e t t e r .  We  they work i n o r d e r t o  s h o u l d r e a l i z e what e f f e c t s our  a c t i v i t i e s have upon t h e i r c o n t i n u e d a b i l i t y t o produce what we  require. 1.4  1.4.1  A Land Use  Perspective  General i n t r o d u c t i o n The  settlement area,  purpose of t h i s s e c t i o n i s t o r e l a t e human and  i n d u s t r i a l development i n the g e n e r a l  t o moose and  their habitat.  study  A l t h o u g h the main  subject